The Early Evolutionary

History of Planktonic Foraminifera

British Micropalaeontological Society Publication Series

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 The Early Evolutionary History
of Planktonic Foraminifera

by
M.K. BouDagher-Fadel,
F.T. Banner
and J .E. Whittaker

with a contribution from

M.D. Simmons

plate digitization by
H. Taylor

lal'l SPRINGER-SCIENCE+BUSINESS MEDIA, B.v.

First edition 1997
1997 Springer Science+Business Media Dordrecht
Originally published by Chapman & Hali in 1997
Softcover reprint ofthe hardcover Ist edition 1997

Typeset in 10/12 Times by Florencetype Ltd, Stoodleigh, Devon

ISBN 978-94-010-6466-8 ISBN 978-94-011-5836-7 (eBook)

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 Contents

List of contributors ix
Preface and acknowledgements xi

1 Introduction
M.K. BouDagher-Fadel, F.T. Banner and l.E. Whittaker
1.1 What are the planktonic foraminifera? 1
1.2 Notes on the nomenclature of morphological structures in taxonomy and time 2
1.3 Notes on the crystallographic structure and chemical composition of the test wall 4

2 The Jurassic Favusellacea, the earliest Globigerinina 17

MD. Simmons, M.K. BouDagher-Fadel, F.T. Banner and l.E. Whittaker
2.1 Origins 17
2.2 The published taxa 20
2.3 Key to the genera 20
2.4 Systematics 20
2.5 Concluding comments 29

3 The Early Cretaceous families 53

M.K. BouDagher-Fadel, F.T. Banner and lE. Whittaker
3.1 Introduction 53
3.2 Key to the suprageneric taxa 54

4 The Favusellidae - the Cretaceous acme of the Favusellacea 57

M.K. BouDagher-FadeL, F.T. Banner and lE. Whittaker
4.1 Introduction 57
4.2 Systematics 59
4.3 Concluding comments 62
5 The practical taxonomy of the Praehedbergellidae 79
M.K. BouDagher-Fadel, F.T. Banner and l.E. Whittaker
5.1 Introduction 79
5.2 Criteria within the species-groups 80
5.3 Supraspecific criteria in the Praehedbergellidae 82
vi Contents

6 The earliest PraehedbergeUidae - Gorbachikella 85

M.K. BouDagher-Fadel, F.T. Banner and 1 E. Whittaker
6.1 Introduction 85
6.2 Key to the species of Gorbachikella 87
6.3 Systematics 87
6.4 Concluding remarks 88
7 The flowering of the PraehedbergeUidae: Praehedbergella 103
M.K. BouDagher-Fadel, F.T. Banner and 1.E. Whittaker
7.1 Introduction 103
7.2 Key to the species of Praehedbergella 105
7.3 Systematics 105
7.4 Concluding comments 110

8 Blefuscuiana, the longest ranging and most diverse of the PraehedbergeUidae 121
M.K. BouDagher-Fadel, F.T. Banner and 1 E. Whittaker
8.1 Introduction 121
8.2 Key to the species of Blefuscuiana 123
8.3 Systematics 124
8.4 Concluding comments 133

9 The PraehedbergeUidae with elongate chambers - Lilliputianella,

Lilliputianelloides gen.nov., and Wondersella 161
MK. BouDagher-Fadel, F.T. Banner and lE. Whittaker
9.1 Introduction 161
9.2 Systematics 163
9.3 Concluding comments 166
10 The planispiraI Schackoinidae (Blowiella, Claviblowiella gen.nov.,
Globigerinelloides, Leupoldina and Schackoina) 177
MK. BouDagher-Fadel, F.T. Banner and lE. Whittaker
10.1 Introduction 177
10.2 Systematics 179
10.3 Concluding remarks 186
11 The evolution of the HedbergeUidae 203
MK. BouDagher-Fadel, F.T. Banner and lE. Whittaker
11.1 Introduction 203
11.2 Systematics 203
11.3 Concluding comments 204

12 The evolution of the PlanomaIinidae 209

M.K. BouDagher-Fadel, F.T. Banner and lE. Whittaker
12.1 Introduction 209
12.2 Systematics 211
12.3 Concluding comments 215
 Contents vii

13 Aspects of the probable physiology and ecology of the Praehedbergellidae,

aided by studies of living Globigerinacea 229
M.K. Boudagher-Fadel, F.T. Banner and lE. Whittaker
13.1 Introduction 229
13.2 Comparison with living morphological analogues 230
13.3 Classification of living Globigerinacea 230
13.4 Conclusions from comparisons with living analogues 235
13.5 Palaeoecology 235

Appendix 1 Planohedbergella gen.nov. 245

M.K. BouDagher-Fadel, F.T. Banner, lE. Whittaker and Lt. D. McCarthy

References 251
Systematic index 259
Subject index 265
 Contributors

M.K. BouDagher-Fadel [Chapter 2]

Department of Geological Sciences M.D. Simmons
University College London Department of Geology and Petroleum Geology
Gower Street University of Aberdeen
London WClE 6BT Meston Building
UK King's College
Aberdeen AB9 2UE
F.T. Banner UK
Department of Geological Sciences
University College London [Plate digitization]
Gower Street H. Taylor
London WClE 6BT Science Photography Unit,
UK The Natural History Museum
and Cromwell Road
Department of Palaeontology London SW7 5BD
The Natural History Museum UK
Cromwell Road
London SW7 5BD [Appendix 1]
UK Ll. D. McCarthy
Department of Geological Sciences
J.E. Whittaker University College London
Department of Palaeontology Gower Street
The Natural History Museum London WCIE 6BT
Cromwell Road UK
London SW7 5BD and
UK Department of Palaeontology
The Natural History Museum
Cromwell Road
London SW7 5BD
UK
 Preface
and Acknowledgements

This book attempts to describe the earliest known Academy of Sciences). Without their assistance in
planktonic foraminifera, of the Jurassic, and to particular, our understanding of the early planktonic
portray their evolution into the widespread and foraminifera would have been much the poorer.
holoplanktonic taxa of the Early Cretaceous. Much of the SEM photography has been
In doing this we estimate we have dealt with specially prepared for the book by two of the
over 135 species, belonging to 26 genera. It has authors (M.K. B-F. and J.E.W.) on the University
been our policy to illustrate, with SEM imagery College, London (Zeiss DSM 940) and The
whenever possible, holotypes or other primary Natural History Museum's (Hitachi S 2500)
types, topotypes and other reliable specimens. To instruments, respectively. Many more illustrations
this end, we have been generously helped by many have been digitized from the original negatives
micropalaeontologists and other technical workers, held in both those institutions from earlier publi-
without whose assistance the book could not have cations of the authors and our collaborators
been prepared. The systematic taxonomy is our (notably Drs Damini Desai, Phil Copestake,
responsibility, however, and we must be blamed Marigold White, Mike Simmons and Tatiana
for any errors which may have occurred. Gorbachik), over the last decade. The remainder
Our fellow enthusiast Dr Mike Simmons have been captured and digitized from a very wide
(University of Aberdeen) is thanked for providing list of publications, the publishers of which (wher-
access to much of the type material of the Jurassic ever reasonably possible) have been contacted and
species, from his many contacts in the former their permission sought to publish; sources of
Soviet Union. We are particularly grateful to him these illustrations are given in the relevant plate
for the many stimulating discussions on the phy- explanations. All the plates have been assembled
logenies and wall composition of these forms. It and digitally produced by Mr Harry Taylor of The
is our pleasure that he has agreed to be the senior Natural History Museum's Photographic Unit
author of Chapter 2. using an Adobe Photoshop programme. In the 64
Professor Tatiana Gorbachik (Moscow State plates which add so much to the value of this book,
University) helped us with both loaning specimens over 1000 negatives and images have been
and valuable discussions. Other specimens were captured in this way and numbered by Harry, the
kindly lent to us by Dr Kirilla Kuznetsova (Moscow ensuing result reflecting his professionalism and
Academy of Sciences), Dr Svetlana Yakovleva care. For these reasons he deserves more than the
(VNIGRI, St Petersburg), Dr G.K. Kasimova customary cursory acknowledgement.
(Azerbaijan Academy of Sciences) and Dr A. We wish to thank Professors AR. Lord and
Babazade (Geological Institute of Azerbaijan G.D. Price (University College London) for their
xii Preface and acknowledgements

enthusiastic help, unfailing support and the use of (United States National Museum of Natural
facilities throughout this study. Dr Paul Bown History, Smithsonian Institution, Washington, DC)
(UCL) kindly used his extensive knowledge of facilitated the loan of Glaessner's metatypes
nannofossils to independently date many of the and of the paratypes of several of Cushman's
rocks which yielded planktonic foraminifera. Dr species. Dr F. Gradstein (now of Saga Petroleum,
Ann Holbourn (now of The Natural History Sandvika, Norway) loaned us figured material
Museum) gave us material from the south-east from Starn (1986) in the collections of the Atlantic
Indian Ocean. Messrs J. Davy and C. Stuart (UCL) Geoscience Centre, Bedford, Canada while Dr M.
are thanked for their technical support, while Dr Bubik (Czech Geological Survey, Brno) sent us
Giles Miller and Greg Welsh (The Natural History SEM images and specimens of Praehedbergellidae
Museum) helped greatly by their expert drafting from the West Carpathians. We sincerely thank
of the range charts and in the finalizing of the them all.
manuscript, respectively. Dr Gordon Cressey and Some of Llynne McCarthy's (University
John Francis (Mineralogy Department, The College, London) PhD study on Late Cretaceous
Natural History Museum) are warmly thanked for planktonic foraminifera Uointly supervised by
their important X-ray analyses of the early plank- F.T.B. and J.E.W.) has impinged on our own
tonic foraminiferal tests, and for their discussions. systematics. It is our pleasure to invite her con-
We are grateful to Dr Geoff Eaton (now of The tribution to the Appendix on Planohedbergella
Natural History Museum) and Dr Stan Duxbury gen.nov.
(now of Duxbury Stratigraphic Consultants) for Finally, we were very grateful for financial
organizing the loan of material from British support from the Natural Environment Research
Petroleum Exploration Co. Ltd and from British Council and the Royal Society (The Daphne
Gas. Dr Phil Copestake (IEDS Ltd, Tetbury, Jackson Memorial Fellowships Trust).
Gloucestershire) was of great value to us in his This has been a labour of love for the co-authors
elucidation of North Sea biostratigraphy. who have spent the last few years trying to unravel
We are also indebted to Dr M.J. Brolsma and the origines) of the planktonic foraminifera and
Dr M. Wannier (Shell International Petroleum Co. elucidate their systematics and phylogenies. We
Ltd, UK) for providing us, specifically for this feel a little like Gulliver in his travels (Jonathan
research, with the Tunisian Early Cretaceous Swift, 1726) in the lands of Blefuscu and Lilliput
samples and to Dr H. Bismuth (SEREPT) for his in that we seem to have seen a very large number
search for, and for sending to us all the published of miniature ovoids; whether they were displayed
research on the Cretaceous of Tunisia. by their bluntly rounded ends or by their pointed
Professor M. Moullade (University of Nice) and ends, perhaps the reader is now a little wiser!
Dr A. Gorog (Eotvos University, Budapest) loaned
us topotypic specimens which we have imaged.
Drs. B.T. Huber, M. Buzas and F.J. Collier London, February 1997
 1
Introduction
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

1.1 WHAT ARE THE PLANKTONIC The planktonic foraminifera (the Globigerinina)
FORAMINIFERA? are always multilocular; there are no single-
chambered planktonic foraminifera. The secreted
Foraminifera are free-living protozoa which grow calcitic or aragonitic tests are always perforated
an elaborate, solid skeleton. The benthonic forms by multitudinous small holes. These perforations,
occur on a wide variety of substrates at all depths which also occur in many benthonic taxa, enable
in the marine realm, whereas the planktonic taxa internal cytoplasm to make biochemical contact
float freely in the upper water of the world ocean. with a cytoplasmic sheet on the outer surface of
Foraminifera vary in size from less than 100 f1m the test (extrathalamous cytoplasm) and with
in diameter to a maximum breadth of many the surrounding sea water. Most importantly, the
centimetres; planktonic species do not exceed planktonic foraminifera lack the additional skeletal
600 f1m in maximum skeletal diameter. Benthonic structures characteristic of benthonic taxa. There
foraminiferal tests may be agglutinated (quartz or are no plugs, no pillars and no canal systems.
other inorganic particles being stuck together by Although the aperture may be modified exteriorly
calcitic or organic cements) or primarily secreted; by the development of apertural lips or plates
these latter may be calcite or aragonite or (rarely) (called portici or tegilla), there are no internal
silica. Planktonic foraminifera have tests made structures in any way comparable to the tooth-
only of secreted calcite or aragonite (see below). plates of benthonic taxa.
Foraminiferal tests rarely consist of only one Foraminiferal tests as a whole have been
chamber; usually, as the organism grows, it adds systematically described (at generic and supra-
successively additional, progressively larger generic level) by Loeblich and Tappan (1964,
chambers, to produce a skeleton of varying 1988), what is known of living foraminifera has
complexity. The benthonic foraminiferal test may been reviewed by Lee and Anderson (1991) and
be made even more complex by the development their colleagues, while the biology of modem
of canal systems within the walls, plugs and pillars planktonic foraminifera has been presented by
within the septa and umbilici, and internal tooth- Hemleben, Spindler and Anderson (1989).
plates modifying the routes of exit and ingress of In this book, we describe what we know of
the cytoplasm through the aperture. the earliest fossil planktonic foraminifera of the
2 Introduction

Jurassic and the Early Cretaceous, previously a did not attempt to investigate the Early Cretaceous
rather neglected field. These microfossils are and older planktonic taxa.
known, of course, only by their tests. The criteria Many attempts have been made to use the
noted above (and in more detail in subsequent Cretaceous planktonic foraminifera for zonal
chapters) have been used to determine that they purposes (e.g. Sigal, 1965; Longoria, 1974; Caron,
were indeed planktonic; while the other micro- 1985, etc.) but in this book we have referred our
fossils associated with them have helped to ascer- taxa mostly to the geostratigraphic age and stage
tain their deep-water marine habitat and, in some in which they occur.
cases, to constrain their age determination. We have endeavoured to make use of a widest
Because of the abundance of the planktonic selection of material from the Jurassic and Early
foraminifera in close stratigraphic sequence Cretaceous marine sediments of the world, thanks
through sediments, and because of the regularity to cooperation of our friends and co-workers
of their structures and their taxonomic diversity, (Acknowledgements) and to make our coverage of
they have provided continuous evidence of evolu- the taxa as comprehensive as possible. From our
tionary changes. These changes were found useful researches of many years this has now lead to a
by both academia and by the hydrocarbon industry thorough revision of the taxonomy, nomenclature
for correlation in sedimentary sequences and for and probable phylogenies of this most interesting
the construction of Late Mesozoic-Cenozoic and at the time, emerging group of foraminifera.
chronostratigraphic scales. In particular, the petro- All our own material, but most notably samples
leum exploration industry found the planktonic from Tunisia, Azerbaijan and the Kacha River of
foraminifera of great importance, because they Crimea were dated for the purpose of this book
were easy to extract from both outcrop and subsur- by nannofossils (and in some cases by ammonites)
face samples and enabled biostratigraphic dating by colleagues at University College London.
to be carried out in new exploration areas very
quickly; examples of their use are the publications
sponsored by Exxon (Stainforth et at., 1975), the 1.2 NOTES ON THE NOMENCLATURE
Royal Dutch/Shell Group (Postuma, 1971) and OF MORPHOLOGICAL STRUCTURES IN
British Petroleum (Blow, 1979). Of these publi- TAXONOMY AND TIME
cations, one (Postuma, 1971) presented illustra-
tions of Albian and younger Globigerinina, while The names which have been given by various
the other two (Stainforth et at., 1975, and Blow, authors, in the past, to superficially similar struc-
1979) dealt solely with Cenozoic taxa. The plank- tures have often been misleading. These names
tonic foraminifera which have been found in have frequently been used following study of only
sediments of 'Middle' Cretaceous and younger age one taxonomic group of planktonic foraminifera,
have been used for over fifty years for worldwide perhaps from only one short interval of geological
biostratigraphy (Bolli, Saunders and Perch- time. We are endeavouring to use a nomenclature
Nielson, 1985). Unfortunately, there have been no for those structures which separate different,
authoritative and comprehensive texts published although morphologically similar, forms which
solely on the Jurassic and Early Cretaceous plank- occur in different families in different geological
tonic foraminifera. The best is the work by periods.
Gorbachik (1986); nothing is comparable to this The perforations which penetrate the walls of
work but it is incomplete, and has been superseded the foraminiferal test, and which allow gas
by the results of more recent investigations. The and solute exchange between intrathalamous and
European Working Group on Planktonic Fora- extrathalamous cytoplasm (Chapter 13), are abun-
minifera, led by Robaszynski and Caron, had dant in the walls of each chamber of the test
produced atlases of the 'Middle' (post-Mid- of all genera of the Globigerinina. They must be
Albian) and Late Cretaceous Globigerinacea distinguished from the pores which occur in the
(Robaszynski, Caron et at., 1979, 1984), but they final chambers of some Cenozoic taxa, which
 Morphological structures in taxonomy and time 3

allow direct ingress and egress of cytoplasmic on the perforation ridges of Parasubbotina
strands (pseudopodia) to and from the test limits. (Olsson et al., 1992, p. 197; Early to Late Palaeo-
Pores occur commonly in the walls of the Miocene cene). True spines really came to full develop-
genera Praeorbulina, Orbulina (Plate 1.4, Fig. 2) ment in the Eocene, and by the end of the Palaeo-
and Globigerinatella, where they replace the gene spinose taxa had replaced muricate forms
primary aperture of ancestral forms; the pores are entirely. Spines are very long, needle-shaped
much larger than the perforations which surround (aciculate) structures, circular or triangular in cross
them. For this reason, the volcano-like structures, section (Hemleben, Spindler and Anderson, 1989,
with axial vents, which were built by extrathal- pp. 209, 210), which have subcylindrical bases
amus cytoplasm over the perforations of many embedded in the chamber wall (Plate 1.4, Fig. 3).
taxa of the Praehedbergellidae are called perfor- In Plate 1.5 note the different distribution of the
ation cones (Plate 1.1, Figs 3, 4), and not 'pore- spine bases found on dead Globigerina (Plate 1.5,
cones' (as they have been called by many authors, Figs 3, 4) and arcadia (Plate 1.5, Figs 5, 6)
especially referring to their occurrence in the from the Recent seas. The sockets for these
Heterohelicacea), because they have nothing to do bases have been imaged in Parasubbotina (op. cit.,
with pores. above). The spine bases which become em-
In the Mesozoic, many Globigerinina have walls bedded in the ridges (Plate 1.4, Figs 1-3) between
with surfaces which possess mound-like structures the macroperforations of post-Palaeogene taxa
sited between the perforations. In the Late have been well illustrated in SEM by Be (1969,
Cretaceous these structures are conical and plates 1-3), Be, Harrison and Lott (1973, plates
pointed, and are often as long or longer than they 1-10), Be and Hemleben (1970, plate 25), and Be
are thick (Plate 1.1, Figs 6, 7); these were named (1980, plates 3, 6 and 7). The spines are well
muricae by Blow (1979, p. 400), and he defined known to aid pseudopodial support, so that the
and discussed them at length (op. cit., pp. 400- pseudopodia can ingest food particles from as far
416; with good illustrations on many of his plates, as possible from the host test, and reject waste
especially plates 208, 210, 211, 221, 223). The matter for similar distances (Hemleben, Spindler
muricae first appear on species of the globigeri- and Anderson, 1989). Clearly, spines have a
nacean genus Hedbergella in the Late Aptian, and greater variety of function than the muricae of the
they can fuse together, along the periphery of the ancestral taxa.
test, to form a muricocarina (in Praeglobo- The Favusellacea of the Jurassic and Early
truncana (Plate 11.1, Figs 5-10) and later Cretaceous (and as stratigraphically young as
Globotruncanidae, as well as in the unique Early Cenomanian) possessed neither pointed
Planomalina (Plate 12.6, Figs 7-13); muricae can muricae nor blunt, scattered pustules. Instead,
also fuse meridionally across the test surface at many of their tests possessed narrow, blunt or
right angles to the periphery, to form a costellae pointed, scattered projections which were more
in Rugoglobigerina (Plate 1.3, Figs 1-6) and elevated and regularly spaced than pustules. We
related genera. Muricae survived in the Globi- call these structures pseudomuricae. Pseudo-
gerinacea to the close of the Eocene. In the muricae are often hollow (Samson et al., 1992;
Cenozoic, several taxa possessed neither muricae Plate 1.1, Fig. I, herein) as muricae can be (Blow,
nor spines, but their surfaces were partly or wholly 1979, plate 208) but never fuse peripherally
covered by irregularly spaced, irregularly rounded into muricocarinae; in contrast, they can fuse into
knobs, which have been called pustules (emended discontinuous ridges (Plate 2.8, Figs 10, 11; Plate
by Blow 1979, pp. 420, 421). These pustules are 2.9, Fig. 11, on Globuligerina) , which are at
very common in genera of the Candeinidae (Plate random angles to each other, and which eventually
1.5, Figs I, 2 and Chapter 13, Plate 13.1, Figs themselves fuse together into favose reticulations
1-6, 11) but are unknown in the Mesozoic. (Plate 2.5, Fig. 11, Favusella), which appear to
At the beginning of the Cenozoic, the sockets be solid (Plate 1.2, Fig. 6). Muricae never fuse
which were once filled with true spines are visible to form reticulations, but occur between the
4 Introduction

macroperforations of the taxa which have them. trocoidea (Gandolfi) and Conoglobigerina mega-
The ridges of Favusella may still retain pseudo- nomica (Kuznetsova) both from the Crimea
muricae at the junctions between the ridges of the (Ukraine), of Globuligerina oxfordiana (Grigelis)
reticulum (Plate 4.4, Fig. 8). Pseudomuricae can from the Russian Plate and Favusella washitensis
enclose patches of microperforations, unlike (Carsey) from Algeria. They found that H.
anything seen in macroperforate forms. trocoidea was calcitic, like all the post-
Therefore, microperforate taxa of the Mesozoic Cenomanian Globigerinina. In contrast, G. oxfor-
can possess pseudomuricae (Favusellacea) or are diana was aragonitic! The specimens of F.
smooth, with no superficial wall structures at all washitensis gave results which were a mixture of
(Globigerinacea, Praehedbergellidae and Schacko- aragonite and calcite, because the aragonitic shells
inidae). In the Cenozoic, microperforate taxa can were partly infilled with secondary calcite. These
possess pustules (GlobasticalGloboconusa of the initial results strongly suggested that the
Danian, and the Candeinidae of the Late Palaeo- Favusellacea, as we shall describe in ensuing
gene to Recent). In contrast, macroperforate taxa chapters, were aragonitic, while the Globigerin-
possess muricae in the Late Cretaceous and acea were calcitic. This would confirm, without
Palaeogene and true spines in many taxa doubt, that our distinction between these two
throughout the Cenozoic. superfamilies (although based primarily on their
gross morphology), would be fully justified.
We have attempted to confirm these initial find-
1.3 NOTES ON THE ings ourselves. X-ray diffraction analyses under-
CRYSTALLOGRAPHIC STRUCTURE taken at The Natural History Museum, London,
AND CHEMICAL COMPOSITION OF using a Gandolfi camera, with 16 to 24 hour
THE TEST WALL exposures of the whole specimen (a non-destruc-
tive technique), were employed. Analyses of both
The hyaline (glassy) foraminifera have a test made Globuligerina oxfordiana (Grigelis) from Russia
of calcium carbonate crystals. The crystals build and France, and G. bathoniana (pazdrowa) from
microgranules (as shown by Banner and Williams, Poland again showed them to be aragonitic. Our
1973) which are themselves aligned into rows topotypic specimens of Favusella washitensis
parallel to the perforations and perpendicular to (Carsey), from Texas, were obviously recrystal-
the surface of the test wall. These crystals are so lized, with an internal filling of secondary calcite
arranged that their optic c-axes are also in align- (Plate 1.2, Fig. 6); they gave, not surprisingly,
ment; when the aligned optic axes are themselves results only for calcite when analysed by our
perpendicular to the test surface, the test is truly method, and is probably due to inversion of the
hyaline. Optically, the crystal structure is not original aragonite to secondary calcite during the
discernible and must be determined by other recrystallization. However, other species we
means. These basic crystallographic criteria, analysed by X-ray diffraction - Conoglobigerina
coupled with the underlying understanding that the dagestanica (Morozova), the type species of
test is of calcite, are fundamental to the definition Conoglobigerina, and C. avarica (Morozova),
of the Globigerinina by Loeblich and Tappan both from Dagestan, and also the topotype of
(1988, p. 452). Moreover, throughout their book, Haeuslerina helvetojurassica (Haeusler), from the
Hemleben et al. (1989) either assume or state that Starn collection - were made of calcite. This,
all Globigerinina are calcitic. again, could be secondary calcite, recrystallized
Analysis of the crystallography and chemical from original aragonite during fossilization, but we
composition of tests of the early planktonic cannot prove this.
foraminifera is still in its infancy. The only work In the best preserved specimens at our disposal
prior to our's was that of Gorbachik and (Globuligerina oxfordiana from the Oxfordian
Kuznetsova (1986). These authors undertook X- of the Moscow region) we were able to under-
ray analysis of specimens of Hedbergella take some high-resolution SEM of fractured
 Structure and chemical composition of the test wall 5

chamber walls (plate 1.2, Figs 2-5). The aragonitic ments soon after deposition (compare the preser-
wall appears to have an innennost layer made with vation of fossil didemnid ascidian spicules, Varol
much smaller microgranules, and the perforations and Houghton, 1996, p. 136). This could explain
appear very thin and sinuous in vertical section. the relative rarity of occurrence of Jurassic
More work is planned on these fascinating and Favusellacean taxa. The crystallography may be
perhaps most fundamental of all criteria. First, we of considerable theoretical value, and it is well
need to find better material from clays and marls known that many benthonic foraminiferal families
(from which our specimens of G. oxfordiana and are consistently calcitic while others are con-
G. bathoniana indeed came) where recrystalliz- sistently aragonitic. This shows that the preferred
ation can be ruled out. mineralogy is genetically controlled, not influ-
From our evidence and that of Gorbachik and enced by the environment they cohabit.
Kuznetsova (1986) we are of the opinion that the If Loeblich and Tappan's scheme (1964, 1988)
Favusellacea were aragonitic, in contrast to the was followed, aragonitic planktonic foraminifera
calcitic Globigerinacea; as yet, however, we do would have to be hived off to a completely separ-
not have the compulsive proof. The instability of ate suborder/superfamily (as indeed are aragonitic
aragonite is well known. Aragonitic fossils are benthic foraminifera). In the meantime, it is far
highly susceptible to dissolution, and their better to emend the definition of the Globigerinina
recovery is unlikely to occur unless there were (as indeed we do in this book) to include both
high sedimentation rates or rapid sealing of sedi- aragonitic and calcitic genera.
Plate 1.1

Figure 1 Globuligerina oxfordiana (Grigelis, 1958). Oxfordian, from the Moscow region, Russia. Enlargement of wall surface
to show pseudomuricae, some apparently hollow (><2250).
Figure 2 Blefuscuiana aptiana (Bartenstein, 1965) s.s. Early Late Aptian, sample 8130, Speeton Cliff, NE England, deposited
in The Natural History Museum, London, no. PF 52084: Enlargement of wall surface to show irregular micro-
perforations (xl400).
Figure 3 Blefuscuiana aptiana (Bartenstein) orientalis BouDagher-Fadel et al., 1996. Early Aptian, sample AGA-l, NE
Azerbaijan, in the neighbourhood of Nardaran village, paratype deposited in The Natural History Museum, London,
no. PF 53017: Showing perforation cones stronger on the early chambers, and microperforations on the last and
penultimate chambers (x230).
Figure 4 Blefuscuiana aptiana (Bartenstein) orientalis BouDagher-Fadel et al., 1996. Aptian, Krasnaya, SW Crimea, Ukraine,
paratype deposited in the Natural History Museum, London, no. PF 53015: Enlargement of wall surface of penulti-
mate chamber showing weak perforation cones (the cones are stronger on the early chambers of the last whorl) (x3200).
Figure 5 Alanlordella bentonensis (Morrow, 1934). Albian, Gault Clay, Arlesey brick pit, S. England, deposited in the Natural
History Museum, London, no. PF 53140: Side view showing macroperforations irregularly scattered over the surfaces
of the chambers, the macroperforations being larger in the early chambers but tending to become much smaller on
the last chamber. This is an evolutionary stage half way between Blowiella and typical Alanlordella (xI75).
Figure 6 Alanlordella banneri BouDagher-Fadel, 1995. Early Albian, SS170, Beauvoir-III Well, Tunisia, holotype deposited in
The Natural History Museum, London, no. PF 53138: Side view showing a macroperforate wall with deep perforation
pits and muricae developed strongly on the early chambers from umbilicus to periphery (x95).
Figure 7 Hedbergella trocoidea (Gandolfi, 1942). Late Aptian, Sirnferopol, Crimea, Ukraine, deposited in the Geological Faculty
of Moscow State University, Russia: Dorsal side showing regularly spaced macroperforations on later chambers and
strong muricae on the early chambers of the last and earlier whorl (xI45).
Plate 1.2

Figures 1-5 Globuligerina oxfordiana (Grigelis, 1958). Oxfordian, from the Moscow region, Russia. Material in The Natural
History Museum, London, showing well preserved aragonitic walls. I, enlargement of the wall surface to show
pseudomuricae, some apparently hollow (x3300); 2, dissection of early whorls to show the inner imperforate mem-
brane and the microperforate aragonitic wall (x2600); 3-5, dissection of later chambers showing the inner lining
becoming perforate, and the canaliculate microperforate aragonitic wall (x2650, x3000, and x5000, respectively).
Figure 6 Favusella washitensis (Carsey, 1926). Early Cenomanian, Shoal Creek, Austin, Texas, USA, topotype deposited
in The Natural History Museum, London: Dissection of (recrystallized) wall, showing canaliculate wall but solid
reticulation (x1300).
Plate 1.3

Figures 1-6 Rugoglohigerina rugosa (Plummer, 1926). Maastrichtian, A. mayaroensis Zone, Guayaguayare/63, Core 19,
Trinidad, deposited in British Petroleum Microfossil Reference Collection at The Natural History Museum, London:
1,3, ventral and dorsal views (xI35); 2, enlargement of surface showing macroperforations and costellae (xl100);
4, 6, ventral and dorsal views showing muricae fused into rows leading from the periphery to the umbilicus
suggesting that the muricae were used both for disaggregation of food particles (very strong muricae on early
chambers) and for directing particles to the umbilical digestive areas (x135); 5, enlargement of the surface showing
muricae fused into discontinuous rows to form costellae (x620).
P1atelA

Figure 1 Hastigerina pelagica (d'Orbigny, 1839). Recent, collected in plankton tow off Bermuda in 0-10 m water depth,
(from Be, 1969, pI. 3, fig. b): enlargement of the perforate wall surface showing triradiate spines (t.s.), the spines
being here triradiate from their bases (xI875).
Figures 2, 3 Orbulina universa d'Orbigny, 1839. Recent, Indian Ocean (live specimens), (from Be, Harrison and Lott, 1973,
pI. 6, figs 4, 5): 2, enlargement of the surface showing pores and originally round spine become triradiate distally
(x780); 3, enlargement of the wall surface showing a spine originating from primary organic membrane, which
divides the inner and outer calcite lamellae (x780), also seen in Figure 2.
Plate 1.5

Figures 1,2 Tenui/ellina/a bradyi (Wiesner, 1931) (= Pylodexia uvula Ehrenberg nomen obli/urn). Recent. In the collections of
The Natural History Museum, London: I, specimen photographed by transmitted light, to compare with SEM
photograph in Figure 2 (x550); 2, showing microperforate, non-spinose test without perforation cones but with
blunt pustules; aperture intraumbilical lacking bulla (x525).
Figures 3,4 Globigerina bulloides d'Orbigny, 1826. Recent. In the collections of The Natural History Museum, London:
3, ventral side of a trochospiral test with subglobular adult chambers and an intraumbilical aperture (xl60);
4, enlargement, showing macroperforate, non-muricate surface, but with abundant and crowded spine bases (x500).
Figures 5,6 Orcadia riedeli (Ragl and Bolli, 1973). Recent. In the collections of The Natural History Museum, London:
5, peripheral view of trochospiral test (x370); 6, detail of the surface showing the macroperforate test with widely
scattered spine bases (xlOOO).
 2
The Jurassic
Favusellacea, the
earliest Globigerinina
M.D. Simmons, M.K. BouDagher-Fadel,
F.T. Banner and J.E. Whittaker

2.1 ORIGINS When such forms reproduce, the progeny are

released into the seawater in the immediate neigh-
The evolution of the earliest planktonic bourhood of the parent test. When a planktonic
foraminifera from their benthonic ancestors is still mode of life is acquired, the chambers lose their
shrouded in uncertainty. adherence and become more fully globular.
As we will describe, the oldest known Planktonic habitats acquired before reproduction
foraminifera which may confidently be considered enable the species to disperse its gametes more
to have been planktonic were species of Cono- freely throughout the seawater and to spread the
globigerina (C. avariformis Kasimova, C. balakh- progeny more widely. The surface tension, at
matovae (Morozova), C. dagestanica Morozova the sea surface, would tend to bring both the breed-
and, possibly, C. avarica Morozova). These ing adults and their released gametes into close
appeared in the Bajocian (Figure 2.2) and are proximity. Many living rotaline genera lose their
known only from eastern Europe (central and benthonic mode of life and become planktonic just
northern Tethys). We consider them to belong to before and during reproduction. The planktonic
the Favusellacea, the aragonitic, microperforate, stage of life adopted by the gamonts of some
pseudomuricate superfamily, with an umbilical species of the Discorbidae and the Cymbalo-
aperture, and all the adult chambers of all of these poridae, for example (Banner et al., 1985), led
species are subglobular in shape. They appear to them to the independent evolution of flotation
show a marked dimorphism, however, which is a structures in sub globular chambers at the last stage
feature not known in true (holo-)planktonic of growth of the test (Plate 2.1). In the Globi-
foraminifera. gerinina, the acquisition of globular chambers
Benthonic, rotaline foraminifera are often resi- enabled the pseudopodia to extrude equally from
dent on firm substrates (stones or algae etc.) and extrathalamous cytoplasm over every part of the
the adherent sides of their tests become flattened. perforate chamber wall (in contrast, an adherent,
18 The Jurassic Favusellacea, the earliest Globigerinina

ventrally flattened chamber would extrude diploid generation would begin with a large
pseudopodia dominantly from the flattened side daughter cell which will form a large proloculus,
only - see, e.g. Alexander and Banner, 1984). to be followed by relatively few chambers per
Some of the Middle to Late Jurassic and whorl - the megalospheric form. The chambers of
Berriasian-Valanginian globigerine foraminifera the earliest whorls of Conoglobigerina spp. are not
(Conoglobigerina) have a restricted geographic always easy to see because of the thickness of the
area; they all occur in south-central and present- chamber wall, but the presence of dimorphic high-
day eastern Europe. They are not recorded from and low-spired forms is often recognizable
anywhere else. The restriction of the area of their (Systematics, below). An exception were the glau-
distribution and the shape of their test seem to conitic moulds of the type specimens of C. conica
indicate a meroplanktonic mode of life (benthonic (Figure 2.1, 4-6) and C. terquemi (Figure 2.1,
in the early stage, becoming planktonic in the 1-3); because they are internal moulds, Iovceva
last stage). The earliest Conoglobigerina (for and Trifonova (1961) could discern the early
example, the Late Bajocian species C. dagestanica chambers easily and drew many chambered
Morozova and C. balakhmatovae (Morozova), (microspheric) early whorls for the high spired
Plates 2.3, 2.4) have early whorls of at least 5 C. conica and relatively few chambers in the
to 6 chambers, while the last whorl has only 4. earliest whorls of the megalospheric, low spired
These early whorls are discorbid-like and were C. terquemi. These two 'species' are strict contem-
probably benthonic, while the last whorls, with poraries (both being found in Tithonian limestones
globular chambers, represent the meroplanktonic near Stubel village, northwest Bulgaria) and as we
stage. have already mentioned, are very likely to be the
This meroplanktonic mode of life seems to have dimorphic pair of a single meroplanktonic species.
been associated with sexual dimorphism, a phys- The youngest Conoglobigerina (Berriasian-Early
ical character unknown in any holoplanktonic taxa.
The early growth stages (the benthonic ones) seem
to have developed in two distinct ways in each
species. One way was to develop a very high spire
with many chambers in the earliest whorls (e.g.
Conoglobigerina conica Iovceva and Trifonova,
Figure 2.1, 4-6); this would be a microscopic
form. The other way was to grow a low dorsal
spire with as many chambers in each whorl as in
the adult; this would be the megalospheric gener-
ation (e.g. Conoglobigerina terquemi Iovceva and
Trifonova, Figure 2.1, 1-3). The origin of this
dimorphism in benthonic foraminifera is clear: the
members of the megalospheric generation were
diploid (had the full set of chromosomes) and
usually reproduced by mitosis (simple replication
of the nuclei), while the microspheric individuals
had been reproduced by meiosis (halving the
number of chromosomes in the parent gametes, Figure 2.1; 1-3 Conoglobigerina terquemi (Iovceva and Tri-
which became haploid). The conjugation of two [onova, 1961). Ventral, peripheral and dorsal views of paratype
gametes would produce a small daughter cell, (from Iovceva and Trifonova, 1961, pI. 2, figs 12-14); x65.
Tithonian, near Stubel village, NW Bulgaria.
which would build a small proloculus, which, in
turn, would be surrounded by many chambers Figure 2.1; ~ Conoglobigeriana conica (Iovceva and
Trifonova, 1961). Dorsal, peripheral and ventral views of holo-
in the first whorl - the microspheric generation. type (from Iovceva and Trifonova, 1961 , pI. 2, figs 2-4); x65 .
In contrast, the mitotically formed succeeding Tithonian, near Stubel village, NW Bulgaria.
 Origins 19

Valanginian) are two fonns described by Praegubkinella, including a new species, P. race-
Gorbachik and Poroshina (1979): C. gulekhensis mosa (Plate 2.2, Figs 1-3, 5), which he (Wernli)
(Plate 2.6, Figs 13-15, high spired, probably believed to be morphologically transitional to
microspheric) and its contemporary C. caucasica Conoglobigerina. All of the species which had
(Plate 2.6, Figs 9-12, low spired, probably been named by Fuchs, as well as most of the
megalospheric); both have the same distinctive fonns figured by Wernli (1995, plates 1,2), appear
chamber shape and wall sculpture. to have strongly concave or flattened ventral
In contrast, the first Globuligerina (G. batho- sides, which were probably the attached sides
niana (Pazdrowa), Bathonian, Plate 2.8, Figs of benthonic fonns (our Plate 2.2, Figs 4, 6, 8,
4-12) has globular chambers throughout ontogeny 9), but P. racemosa Wernli (Plate 2.2, Figs 1-3,
and was holoplanktonic; it is known through 5) does appear to have been free-living, probably
eastern and western Europe and to Canada (Stam, with an umbilicate but otherwise convex
1986). The taxon, like its descendant, G. oxford- ventral side. It may well have been ancestral, in
iana (Grigelis), therefore, no longer had a Toarcian time, to the earliest (Bajocian)
restricted geographic distribution. Conoglobigerina.
The origins of this evolutionary story have Fuchs (1975) may have been right in one sense,
been proposed in a series of papers published by in that an oberhauserellid (if not Oberhauserella
Fuchs between 1964 and 1979. Fuchs described itself) did give rise to Conoglobigerina; it would
over 20 species, most of them being new, from also be attractive for the aragonitic Oberhauser-
the Triassic of Austria and Jurassic of Poland. He ellidae to be one possible ancestor of the (aragon-
referred to the Triassic fonns as 'Globigerina-like' itic) Favusellacea. On the other hand, his
and he thought them to be planktonic. However, evolutionary hypothesis that Mariannenina was the
many research workers have subsequently ancestor of Hedbergella is not tenable. Marianne-
examined Fuchs' material (Drs A. Gorog of nina Fuchs (1973) is a genus of uncertain status ('it
Hungary, and F. Rogi of Austria, pers. comm.) may be congeneric with Jurassorotalia but it needs
and have concluded that it consists mainly of study of better preserved specimens' , according to
badly preserved and recrystallized benthonic Loeblich and Tappan, 1988, p. 705). Furthennore,
specimens. Examination of his figures strongly it seems to have a double peripheral keel, a char-
suggests that most of his taxa are indeed acter absent in Jurassic or even Early Cretaceous
benthonic; the ventral side is flat, the umbilicus is planktonic foraminifera.
closed and the specimens show no evidence of Therefore, we conclude that Fuchs' Triassic
planktonic habitat at all. According to Fuchs' 'globigerinids' are benthonic fonns and Cono-
(1975) scheme, his Triassic genus Oberhauserella globigerina evolved in the Jurassic with the first
was the direct ancestor of the Jurassic genus cycle of zooplanktonic life. However, the Toarcian
Conoglobigerina, and that the Triassic Schmidita sediments of Europe did yield one species of
Fuchs was the ancestor of the Callovian Praegubkinella which may have been the first true
Mariannenina; he believed that this last genus ancestor to the meroplanktonic Conoglobigerina.
evolved, in tum, into the holoplanktonic genus Subsequently, the Favusellacea became holo-
Hedbergella. However, according to Loeblich planktonic (fully planktonic) and widespread with
and Tappan (1988, p. 439), Oberhauserella and the first Globuligerina which appeared in the
Schmidita were benthonic fonns and they both Bathonian. Much more work needs to be done
belong to the benthic family Oberhauserellidae, in before a clearer picture can emerge.
the aragonitic Robertinina. There are no known species of Conoglobigerina
More recently, Wernli (1995) has described in sediments older than Late Bajocian, but, as so
from the Toarcian (Early Jurassic) of Switzerland many species occur there, it is probable that their
specimens that he has referred to as Oberhauser- sudden appearance may relate to rising eustatic
ella quadrilobata Fuchs (1967) (reproduced in sea-level in the Bajocian (Haq et al., 1988),
Plate 2.2, Figs 4, 7, 9, herein) and to Fuchs' genus opening up new niches.
20 The Jurassic Favusellacea, the earliest Globigerinina

2.2 THE PUBLISHED TAXA 2.3 KEY TO THE GENERA

With the exception of the species of Globuli- 1 Aperture intraumbilical

gerina, the Jurassic taxa are poorly known; 1.1 aperture an interiomarginal, low arch
published descriptions and names are based upon Conoglobigerina
very few specimens, and the original illustrations 1.2 aperture loop-shaped and high
are often unreliable, crude drawings. However, the Globuligerina
situation has been greatly improved by the work 2 Aperture intra-extraumbilical
of Gorbachik, Simmons and their colleagues (in 2.1 chambers inflated, intercameral sutures
press), as many of the published taxa (mainly from depressed Haeuslerina gen.nov.
the former Soviet Union) have been re-imaged by 2.2 chambers not inflated, intercameral sutures
SEM for the first time - primary types and topo- not depressed
types can now be recognized. Thanks to the coop- Compactogerina gen.nov.
eration and enthusiasm of Professor T. Gorbachik
and Dr K. Kuznetsova, in particular, these speci-
mens have been specially photographed by one of 2A SYSTEMATICS
us (J.E.W.) at The Natural History Museum,
London. The resulting plates (plates 2.3-2.9) Suborder GLOBIGERININA Delage and
contain virtually every species of planktonic Herouard, 1896
foraminifera so far described from the Jurassic and
earliest Cretaceous and because of their impor- Superfamily FAVUSELLACEA Longoria, 1974
tance are reproduced here. In spite of this, it must (nom. corr.) emend. Banner and Desai, 1988
be remembered that, in the main, the taxa are still
represented only by few specimens, and their Family CONOGLOBIGERINIDAE Jam.nov.
intraspecific variation and stratigraphic ranges are
still imperfectly known. In the absence of conclu-
Diagnosis: Those genera of the Favusellacea
sive evidence of dimorphism we will treat, in this which do not possess a favose surface structure of
chapter, the potential dimorphs as separate taxa. fused pseudomuricae forming an anastomosing
Other species names have been proposed for reticulation.
tests which were believed by the authors to be of
Jurassic age, but which are in fact contaminants! Genus Conoglobigerina Morozova, 1961 emend
These are illustrated on Plate 2.10. Their surface Type species Globigerina (Conoglobigerina)
morphology shows them all to be of Cenozoic age. dagestanica Morozova, 1961
They are: Globigerina gaurdakensis Balakhmatova
and Morozova, 1961 (Plate 2.10, Figs 8-11), orig-
Emended diagnosis: Conoglobigerinidae with an
inally claimed to be Late Bajocian; Globigerina
intraumbilical aperture, which is an interiomarginal
spuriensis Bars and Ohm, 1968 (Plate 2.10, Figs
low arch. [Synonym: Woletzina Fuchs, 1973].
1-7), originally thought to be Late Bajocian-Early
Bathonian, but similar to several Palaeogene Conoglobigerina dagestanica Morozova, 1961
species (the holotype - Plate 2.10, Figs 4-7 - may
(Plate 2.3, Figs 1-13; Plate 2.4, Figs 5, 13-15;
be a Catapsydrax); and Globuligerina geczyi
Figure 2.2)
Gorog, 1994 (Plate 2.10, Figs 12-14), said to be
Early Jurassic but recognised by us to be Danian 1961 Globigerina (Conoglobigerina) dages-
(Plate 2.10, Figs 12,14, is a Praemurica sp.; Plate tanica Morozova (in Morozova and
2.10, Fig. 13, the holotype, is Globoconusa daub- Moskalenko), p. 26, pI. 1, figs 13-15 (holo-
jergensis (Bronnimann. These 'species' are dis- type), pI. 2, figs 14-19, text-figs 7; 1-24.
cussed in more detail by Gorbachik et al. (in press). 1961 Globigerina (Eoglobigerina) balakh-
Nevertheless, their names must now be rejected. matovae Morozova (pars) (in Morozova
 ... ... ... ... ... ... ...
~ IS ~ I
II> : :,. ~ :... e ~
VOLGIAN
BAJOCIAN BATHONIAN CALLOVIAN OXFORDIAN KIMMERIDGIAN TITHONIAN VALANGINIAN
BERRIASIAN

EARLY LATE EARLY LATE EARLY LATE EARLY LATE EARLY LATE EARLY LATE EARLY LATE EARLY LATE ~
-- - Conogloblgerlns svsrlca
C. svsrllonn/e
- C. bBlskhmstovse
C. CBUcaSIeB
C. conics
C. dsgestsnlcB
C. guleklHlnsls
C. /Ursssica
C. megsnomleB
-- C. terqutJml

~ Globullgerins oxfordlBns
G. bBthonlBns
G. callovlansls
HBeus/erins hetveto/UnssleB
H.psrvB
- CompBctogerins stellBpolBrls

Figure 2.2 Stratigraphical distribution of the Conoglobigerinidae (Favusellacea) - Conoglobigerina, Globuligerina, Haeuslerina gen.nov. and Compactogerina gen.nov.
Chronostratigraphy and Absolute Time Scale after Gradstein et al. (1995).
22 The Jurassic Favusellacea, the earliest Globigerinina

and Moskalenko), p. 23, pI. 1, figs 9-11 chambers are inflated and subglobular, with
(paratype) only. depressed intercameral sutures. The surface of the
1961 Globigerina (Conoglobigerina) avarica wall is covered with pseudomuricae (Plate 2.3,
Morozova (pars) (in Morozova and Fig. 11), which often fuse laterally into very short
Moskalenko), text-fig. 8; 8,9, 15, 17, 18 ridges. The aperture is a low intraumbilical arch
(paratypes) only. with no known lip; there are suggestions that
1980 Conoglobigerina dagestanica Morozova; a small bulla may occasionally be present (Plate
Grigelis and Gorbachik, pI. 1, figs 1, 2. 2.3, Fig. 10).
1984 Globuligerina araksi Kasimova (in Kasi- It is possible that C. dagestanica was dimorphic,
mova and Ayileva), pp. 12, 13, figs lOa-c. the microspheric generation (like the holotype
1986 Conoglobigerina dagestanica Morozova; illustrated by Morozova, and the paratypes figured
Gorbachik, pI. 1, figs 1-3. by us, Plate 2.3) having five to six chambers per
whorl initially, but the megalospheric generation
Remarks: Morozova (1961) obtained her type (like that figured by Morozova, 1961, plate 2, Fig.
specimens from the Bathonian of central Dagestan. 17) having only four. The height of the spire, in
Globuligerina araksi was described by Kasimova each case, was much the same. It does not seem
(in Kasimova and Ayileva, 1984) from the Upper possible to distinguish microspheric and megalo-
Bajocian of Azerbaijan. We consider them spheric generations by the height of the spire
synonymous. Both (Plate 2.3, Figs 1-13) are high alone. It was suggested by Banner and Desai
spired, with four chambers in the final whorl. (1988, p. 147) that dimorphism in Conoglobi-
Morozova (1961) believed the first whorl of the gerina indicated that the genus was meroplank-
test to have five chambers, with this number tonic (at least in the microspheric generation) and
reducing to four or even three in the last formed therefore close to the benthonic ancestor of the
whorl. Morozova (op. cit.) figured many paratypes Favusellacea, but this thesis is not yet proven.
by photography (1961, pI. 2, figs 14-19) and line
drawings (text-figs 7; 4-24); like the paratypes Range: Late Bajocian to Bathonian. This species
illustrated in this book (Plate 2.3, Figs 8-13), all was found by Morozova (1961) in the Early
have four chambers in the last whorl. The only Bathonian and Late Bajocian of Dagestan and
specimen illustrated by Morozova (1961, op. cit.) Turkmenia, and in the Late Bajocian of Azerbaijan
as having three chambers in the last whorl was by Kasirnova and Ayileva (1984).
the drawing of the umbilical surface of the holo-
type (text-fig. 7; 3); however, our new photograph Conoglobigerina avarica Morozova, 1961
of the same surface of the holotype (Plate 2.3, Fig. (Plate 2.4, Figs 1-4, 6; Figure 2.2)
7) shows it to have four chambers in the last whorl,
just like the paratypes. Therefore, we conclude that 1961 Globigerina (Conoglobigerina) avarica
there is no evidence for a primary type of this Morozova (in Morozova and Moskalenko)
species possessing only three chambers in the last (pars), p. 28, pI. 2, figs 1-8 (1-4 holotype),
whorl. 10-13,20; text-figs 8; 1-6, 13, 14, 19-21
Rare paratypes of C. avarica Morozova (Plate (?7,?11, ?12; non 8, 9,15,17, 18).
2.4, Fig. 5) and C. balakhmatovae (Morozova) 1971 'GZobigerina' avarica Morozova; Bronni-
(Plate 2.4, Figs 13-15) are referable to C. dages- mann and Wernli, p. 123, pI. 2, figs 1-4,
tanica. They all have four chambers abutting the pI. 4, figs 3, 4.
umbilicus, and spires which are as high as in
some paratypes of C. dagestanica. Remarks: C. avarica is a very high spired
C. dagestanica (and, of course, its synonym C. species, and the spire of the holotype itself is fifty
araksi) has a high spire with at least five cham- per cent higher than the test is broad (our new
bers in each of the earliest whorls (at least in both photograph of the holotype (Plate 2.4, Fig. 1)
holotypes), reducing to four in the final whorl. The shows this well). The same ratio is seen in our
 Systematics 23

figured paratype (Plate 2.4, Fig. 1). There are three C. avariformis is close to C. dagestanica but
subglobular chambers in the final whorl; the aper- possesses much more depressed chambers, less
ture is an arch within the small umbilicus. There depressed, more oblique intercameral sutures, and
are about five chambers in the earliest whorls, a more compact test.
judging from the paratypes figured here (Plate 2.4,
Fig. 6) and by Morozova (1961, pI. 2, fig. 12). Range: Late Bajocian. This species is known
The paratypes figured by Morozova (1961) with only from the Late Bajocian of Azerbaijan
four chambers in the last whorl (including that (Nakhichevan).
figured here, Plate 2.4, Fig. 5) are referred by us
to C. dagestanica. The wall of the test may seem Conoglobigerina balakhmatovae (Morozova,
smooth under the lower magnification, but SEM 1961)
shows it to be covered by pseudomuricae (Plate (Plate 2.4, Figs 7-12, 16; Figure 2.2)
2.4, Fig. 4).
1953 Globigerina sp. Balakhmatova, p. 88, text-
Range: ?Late Bajocian-Early Bathonian. The figs la-c.
type specimens were obtained by Morozova from 1961 Globigerina (Eoglobigerina) balakhma-
the Early Bathonian of central Dagestan (Morozova tovae Morozova (in Morozova and
in Morozova and Moskalenko, 1961). Bronnimann Moskalenko) (pars), p. 23, pI. 1, figs 1-8,
and Wernli (1971) recorded their specimens from 12 (not figs 9-11).
the Middle Jurassic ('Dogger') of the southern Jura, 1971 'Globigerina' balakhmatovae Morozova;
France, which is also Bathonian in age but may Bronnimann and Wernli, p. 121, pI. 1, figs
range into the Late Bajocian as well. 1-4, pI. 4, figs 1,2.
1986 Globuligerina balakhmatovae (Morozova),
Conoglobigerina avariformis Kasimova, 1984 Gorbachik, pI. 4, Figs 1-3.
(plate 2.7, Figs 8-10; Plate 2.8, Fig. 2; Figure 2.2)
Remarks: The test is low spired and the dorsal
1984 Conoglobigerina avariformis Kasimova surface is only slightly convex. This species also
(in Kasimova and Aliyeva), p. 2, pI. 1, figs differs from C. dagestanica Morozova in having
Ia-c, 2-4. a faster rate of chamber enlargement. In C. balakh-
matovae, the hlb =0.7 (approx.), whereas in C.
Remarks: Kasimova figured the holotype of the dagestanica the hlb = 0.5 (approx.). Ventrally,
species as possessing five chambers initially, four chambers abut the umbilicus in each of these
reducing to four chambers in the penultimate species, but about six chambers occupy the first
whorl and then to three in the final whorl. The whorl in C. balakhmatovae (Plate 2.4, Figs 9, 16
metatype illustrated by us (Plate 2.7, Figs 8-10) herein; also Morozova, 1961, plate 1, fig. 1).
may be an immature specimen, with only four Morozova (in Morozova and Moskalenko,
chambers in its last whorl. The umbilical views 1961, p. 61) believed C. balakhmatovae to be
drawn by Kasimova depict an umbilical covering, dimorphic, the microspheric generation having
in each of the four cases illustrated, which might five or six chambers in the first whorl, but the
be a bulla; our metatype (plate 2.7, Fig. 10) may megalospheric possessing only three or four cham-
also have a bulla. The true chambers of the species bers per whorl initially. Her type specimens were
are very depressed and the intercameral sutural from Early Bathonian of Dagestan. The specimen
depressions at the periphery are very broad. from the Bajocian of the north west Caucasus,
The pseudomuricae of the surface may fuse into imaged in SEM by Gorbachik (1986), shows five
very short, discontinuous ridges (Plate 2.8, Fig. 2). chambers per whorl visible initially; Morozova
These features are also found on C. dagestanica; (1961 op. cit.) obtained similar specimens from
they are the stratigraphically earliest examples the Late Bajocian of Turkmenia. Therefore, this
known to us of pseudomurical ridges. generation with five to six chambers initially
24 The Jurassic FavuseUacea, the earliest Globigerinina

ranged from Late Bajocian to Early Bathonian. discontinuous, blunt ridges (Plate 2.6, Fig. 12).
The forms with three or four chambers in the Ultimate elongation of these ridges, and their
earliest whorls appear to have been rare; her illus- fusion to form reticulate, favose patterns, will lead
trations which do not clearly exhibit the number to the evolution of the genus Favusella in the
of chambers in the early whorls may have been following geostratigraphic stage (Hauterivian).
interpreted as possessing few chambers per whorl,
but the whorls may have been obscured by Range: Early Berriasian to Early Valanginian of
sediment. Certainly, there are no specimens the southern Caucasus, Azerbaijan and the eastern
photographed either by Morozova (1961), or by Crimea.
us (plate 2.4) which show so few chambers in the
Conoglobigerina conica (lovceva and Trifonova,
early whorls. In our opinion, dimorphism in this
1961)
species is not yet proven. Nevertheless, it is prob-
able that dimorphism did occur in some species (Figure 2.1; 4-6; Figure 2.2)
of Conoglobigerina, as in C. dagestanica (above).
1961 Globigerina conica Iovceva and Trifonova,
pp. 343-347, pI. 2, figs 1-8.
Range: Late Bajocian to Early Bathonian of
Dagestan, Turkmenia, NW Caucasus and the Remarks: We were unable to borrow any
southern Jura of France. primary type or topotype specimens of this
ConoglobigerilJll caucasica (Gorbachik and species, which Masters (1977, p. 464) claims to
Poroshina, 1979) have been based on glauconitic moulds. Dr
Trifonova kindly sent us unpublished photographs
(Plate 2.6, Figs 9-12; Figure 2.2) of some primary types to supplement the draw-
1979 Globuligerina caucasica Gorbachik and ings which were originally published (Iovceva and
Poroshina, pp. 26-28, text-figs 2a-3d. Trifonova, 1961), some of which are reproduced
here (Figure 2.1; 4-6). The species appears to
possess an arched, intraumbilical aperture,
Remarks: The test has at least five chambers in
conoglobigerine in type, inflated and subglobular
the early whorls, but only four subglobular cham-
chambers, and a very high spire (dlt =1.0--1.3
bers in the last whorl. The height of the dorsal spire
approx). Four chambers comprise the last whorl,
is low, but the dorsal side is clearly convex. The
but many more (seven!) were drawn by the authors
intercameral sutures are distinctly depressed, and
to form the first whorl of the holotype (if that
form intercameral sutural angles of about 130 at
specimen was a glauconitic mould, these earliest
the periphery. The umbilicus is small but open and
chamber moulds may well have been clearly
deep, and the aperture is intraumbilical but (on the
visible); if this is so, the taxon might have been
paratype figured by Gorbachik and Poroshina, 1979,
the microspheric partner of its contemporary ,
text-fig. 3c) no broader than the narrow umbilicus;
C. terquemi (Iovceva and Trifonova) (q.v.) (as
the metatype figured in this chapter, Plate 2.6, Fig.
suggested by Banner and Desai, 1988, p. 147).
11, probably like the holotype illustrated in the
original publication, unfortunately has sediment
Range: Known only from the Tithonian of
covering its umbilicus. The aperture is, however,
north-west Bulgaria.
shown (op.cit., text-fig. 3c) to be conoglobigerine
and not globuligerine. This species must be one Conoglobigerina gukkhensis (Gorbachik and
of the stratigraphically youngest known species of Poroshina, 1979)
Conoglobigerina, and is noticeable for the lack
(Plate 2.6, Figs 13-15; Figure 2.2)
of height of the dorsal spire (in profile, the test of
C. caucasica is very like that of early Favusella). 1979 Globuligerina gulekhensis Gorbachik and
The surface of the test is covered with pseudo- Poroshina, pp. 24-26, pI. 1, figs la-ld,
muricae which occasionally fuse into very short, 2a-2d, text-figs la--c.
 Systematics 25

1983 Globuligerina gulekhensis Gorbachik and 1958 Globigerina jurassica, Hofman, pp. 125,
Poroshina; Gorbachik and Kuznetsova, pp. 126, text-figs a-c.
459-466, pI. 2, fig. 5. 71973 Woletzina jurassica (Hofman), Fuchs, pp.
1986 Globuligerina gulekhensis Gorbachik and 460,461,463, pI. 2, fig. 7, pI. 5, fig. 4.
Poroshina; Gorbachik, p. 80, pI. 10, fig. 3. 1977 Globigerina jurassica Hofman; Masters,
1988 Conoglobigerina gulekhensis (Gorbachik pp. 463, 464.
and Poroshina), Banner and Desai, p. 148. 1982 Woletzina jurassica (Hofman), Banner,
1995 Conoglobigerina gulekhensis (Gorbachik p. 150, Figs 5.4a-c.
and Poroshina), BouDagher-Fadel et al.,
pp. 184-186, pI. 1, figs la, b. Remarks: C. jurassica was designated as the
type and only species of his new genus Woletzina
Remarks: The aperture of this species is not by Fuchs (1973); his drawings are only doubtfully
clear on the metatype figured here (Plate 2.6, of the same species as that described by Hofman
Fig. 15) but there is an indication of a sediment- (1958), however. Hofman obtained her specimens
filled intraumbilical arch with a thin rim. This from the Bathonian-Early Callovian of SE Crimea
aperture and umbilicus are very similar to those (Ukraine). The holotype of this species is lost
drawn by Gorbachik and Poroshina (1979, text- (personal communication, Dr Tatiana Gorbachik),
fig. 1b), and are clearly conoglobigerine. The so we formally designate the paratype figured here
sculpture of the wall surface consists of separate (Plate 2.7, Figs 11-13) as the neotype, following
pseudomuricae and short ridges formed by their article 75 of the ICZN (1985).
fusion; these ridges form broken cells, very close The neotype is high spired with four chambers
to a reticulate, favose pattern, but as yet incom- in the last whorl and an intraumbilical aperture
plete. There can be no doubt that C. gulekhensis covered by a bulla (which has at least two acces-
is very closely ancestral to the earliest Favusella, sory apertures). The surface is covered with
a genus which appears in the Hauterivian (or even pseudomuricae which occasionally fuse into very
slightly earlier, in the Berriasian, according to short, disconnected ridges (Plate 2.8, Fig. 3). C.
Wernli et al., 1995; our Plate 4.2). dagestanica Morozova (the type species of
The chamber shape, umbilical size and shape, Conoglobigerina) also has a bulla (Plate 2.3, Fig.
aperture and wall sculpture are very similar to 10) and as noted above in our introductory section
those of the contemporary C. caucasica (Gorba- ('Origins') bullae are known from the first to last
chik and Poroshina). The principal difference Favusellacea. We have no hesitation in consid-
between the two taxa is the height of the spire ering Globigerina jurassica to be referable to
and very convex dorsal side of the test of Conoglobigerina.
C. gulekhensis. It is possible that C. gulekhensis The specimens drawn by Fuchs (1973) came
is the rnicrospheric form, and C. caucasica the from the Late Callovian and Early Oxfordian of
megalospheric partner of a dimorphic pair (as Poland and the intercameral sutures are more
suggested by Gorbachik and Poroshina, 1979, deeply depressed than those of the neotype; their
p. 24), but the early whorls of each 'species' so identity is not yet proven.
far imaged by SEM are obscure and this relation-
ship is not yet proven. Range: This species is known with confidence
only in the Bathonian-Early Callovian of the
Range: Early Berriasian-Early Valanginian of Ukraine (SE Crimea).
south eastern Caucasus, eastern Crimea and
Azerbaijan. Conoglobigerina meganomica (Kuznetsova,
1980)
Conoglobigerina jurassica (Hofman, 1958)
(Plate 2.5, Fig. 1-3; Figure 2.2)
(Plate 2.7, Figs 11-14, Plate 2.8, Fig. 3; Figure
2.2) 1980 Globuligerina meganomica Kuznetsova (in
26 The Jurassic FavuseUacea, the earliest Globigerinina

Kuznetsova and Gorbachik), p. 750, Figs 1988, p. 147) that C. terquemi is the megalo-
2,5-8. spheric partner of its contemporary C. conica. If
this is so, these taxa would comprise the strati-
Remarks: The aperture and umbilicus on the graphically youngest example of dimorphism in
paratypes figured by the original author (1980, Fig. meroplanktonic Conoglobigerina.
2, 6-8) are obscure, but the SEM photograph of
the paratype illustrated by us (Plate 2.5, Fig. 3) Range: Known only from the Tithonian of
appears to have an aperture of conoglobigerine northwest Bulgaria.
type - an intraumbilical low arch, similar to that
Genus Globuligerina Bignot and Guyader, 1971,
of C. dagestanica Morozova. However, the holo-
emend.
type was drawn by Kuznetsova (1980, Fig. 2,
5a--c) showing the aperture to be an interiomar- Type species Globigerina oxfordiana Grigelis,
ginallow arch, intra-extraumbilical in extent; this 1958
may have been a deformed specimen and should
be re-examined. Emended diagnosis: Conoglobigerinidae with
an intraumbilical aperture which is loop-shaped,
Range: This species was described from the late with a distinct rim.
Early to early Late Callovian of northern Crimea. Globuligerina oxfordiana (Grigelis, 1958)
n is not yet known from elsewhere.
(Plate 1.1, Fig. 1, Plate 1.2, Figs 1-5, Plate 2.9,
Conoglobigerina terquemi (Iovceva and Figs 1-15; Figure 2.2)
Trifonova, 1961)
1958 Globigerina oxfordiana Grigelis, pp. 109,
(Figure 2.1, 1-3; Figure 2.2) 110, figs la--c.
1961 Globigerina terquemi Iovceva and 1971 Globuligerina oxfordiana (Grigelis),
Trifonova, pp. 344-347, pI. 2, figs 9-14. Bignot and Guyader, p. 83, pI. 1, figs 1-3;
pI. 2, fig. 3.
1986 Globuligerina oxfordiana (Grigelis), Gor-
Remarks: The specimens which formed the
bachik, p. 79, pI. 7, figs 1-3; pI. 9, fig. 1.
primary types of this species were found by its
1986 Globuligerina oxfordiana (Grigelis), Starn
authors to occur with the primary types of C.
(pars), p. 105, pI. 7, figs 1-5 (not figs
conica (q.v.), and it is likely that all the specimens
6-12), pI. 8, figs 4-12 (not figs 1-3),
were glauconitic moulds. Like C. conica, the
pI. 9, figs 1-5, pI. 14, figs 3,4,8-15 only.
chambers were drawn (op. cit., above) to be
1986 Globuligerina oxfordiana (Grigelis),
subglobular, with four abutting the umbilicus; the
Wernli and Kindler, pp. 137-147, pI. 1,
aperture is an intraumbilical arch, as broad as
figs 1-4, 17-19.
the narrow umbilicus itself. It is possible that
1987 Globuligerina oxfordiana (Grigelis),
the original authors' drawings show a bulla on the
Riegraf, p. 192, pI. 2, figs 1-7.
holotype (1961, pI. 2, fig. 10) and the remains of
1988 Globuligerina oxfordiana (Grigelis),
a bulla on the figured paratype (1961, pI. 2, fig.
Banner and Desai, p. 146, pI. 1, figs 1-3.
13; reproduced here as our Figure 2.1; 1).
1992 Globuligerina oxfordiana (Grigelis),
However, the spire is very low (height/diameter
Samson et ai., pp. 419, 420, pI. 4, figs 1-14.
ratio of 0.6, compared to the height/diameter ratio
1995 Globuligerina oxfordiana (Grigelis),
for C. conica of about 1.2), and only five cham-
BouDagher-Fadel et ai., p. 182, pI. 1, figs
bers were drawn to comprise the first whorl of
2,3.
both the holotype and the figured paratype. As
remarked above (c. conica), the early whorls
would be clearly visible in a galuconitic mold. It Remarks: This species has been well described
is possible (as suggested by Banner and Desai, by many authors, and its aperture is very variable
 Systematics 27

in shape but is always a bulimine loop, with its 1980 Conoglobigerina bathoniana (Pazdrowa),
posterior margin set forwards into the umbilicus. Grigelis and Gorbachik, pI. 8, fig. 3.
A bulla covers the primary aperture and the 1984 Globuligerina oxfordiana (Grigelis),
umbilicus (Plate 2.9, Fig. 8); if the bulla is broken Bignot and Janin, pp. 755, 756, pI. 1, figs
off, its trace almost always remains. The bulla has 3,6, ?8.
fewer pseudomuricae than the surface of the cham- 1986 Globuligerina bathoniana (Pazdrowa),
bers but these pseudomuricae often fuse into very Starn, p. 108, pI. 9, figs 6-13; pI. 10, figs
short, discontinuous ridges (BouDagher-Fadel et 1-12; pI. 11, figs 1-10; pI. 12, figs 9-12;
aI., 1995, plate 1, fig. 3c) as they do on the surface pI. 13, figs 1-12; pI. 14, figs 1,2,5-7.
of the chambers (Plate 2.9, Fig. 11). There are 1987 Globuligerina bathoniana (Pazdrowa),
about five chambers in each early whorl, reducing Riegraf, p. 195, pI. 1, figs 1-11, Fig. 5a-e.
to four in the last whorl; the H:D ratio can be
quite variable maybe suggesting the possibility of Remarks: This species is very similar to G.
dimorphism. oxfordiana but it has a much higher dorsal side,
The material illustrated here is beautifully and its loop shaped aperture is much lower in
preserved. On analysis by X-ray diffraction the the apertural face. Traces of a bulla may be seen
test proved to be aragonitic (Chapter 1). SEM on many specimens (Plate 2.8, Figs 6, 9) but it
photographs of fractured walls are shown in Plate often appears to have been absent. The pseudo-
1.2, Figs 1-5. muricae are usually separate, but adjacent pairs
could fuse to form very short ridges (Plate 2.8,
Range: This species was originally described Figs 10, 11).
from the Early Oxfordian of Lithuania. It has also
the broadest geographic distribution of any Range: This species was described by Pazdrowa
Jurassic planktonic foraminifera, being known (1969) from Bathonian sediments of Poland; we
from as far afield as Le Havre, northern France illustrate here well preserved topotype material
(Plate 2.9, Figs 5-7,9-11), near Moscow and the (Plate 2.8, figs 4-12). This species has been found
Upper Volga Basin, Russia (Plate 2.9, Figs 8, in sediments ranging in age from Late Bathonian
12-15), the Grand Banks of Newfoundland and to Early Kimmeridgian in eastern Canada,
Ethiopia. All of the occurrences are Oxfordian in Portugal, northern France, northern and southern
age (probably Early Oxfordian). Germany, northern Switzerland, southern Poland,
Azerbaijan and the Crimea. Some records as early
Globuligerina bathoniana (Pazdrowa, 1969)
as Late Bajocian have yet to be proven, as are
(Plate 2.8, Figs 4-12; Figure 2.2) those of Starn (1986) and Riegraf (1987) up into
the Early Valanginian.
1968 Globigerina helvetojurassica Haeusler;
Oesterle, pp. 774-778, text-fig. 50 (non Globuligerina calloviensis Kuznetsova, 1980
Haeusler, 1881). (Plate 2.5, Figs 5-10; Figure 2.2)
1969 Globigerina bathoniana pazdrowa, pp. 45,
46, pI. 2, figs 1-9; pI. 3, figs 1-3; pI. 4, 1980 Globuligerina calloviensis Kuznetsova (in
figs 1-3; text-figs 1-9. Kuznetsova and Uspenskaya), p. 750, pI.
1973 Conoglobigerina bathoniana (Pazdrowa), 2, figs 1-4.
Fuchs, p. 455, pI. 1, fig. 2. 1986 Globuligerina calloviensis Kuznetsova;
1976 Gubkinella bathoniana (Pazdrowa), Gorbachik, pI. 5, fig. 3, pI. 6, figs 1,2.
Ascoli, pI. 1, figs la, b.
1976 'Globigerina' bathoniana Pazdrowa; Remarks: The original specimens figured by
Gradstein, pI. 1, figs 5, 6. Kuznetsova (1980, as drawings and SEM images)
1977 Globigerina hoterivica Subbotina; were reasonably well preserved but their umbili-
Masters, p. 460, pI. 22, figs 1-3. cal characters were not clear. However, the
28 The Jurassic Favusellacea, the earliest Globigerinina

metatype which we figure here (Plate 2.5, Figs level for his new species was the zone of
5-7) and a topotype illustrated by SEM by 'Ammonites transversarius' in the Birmensdorfer
Gorbachik (1986, plate 6, fig. 2) show the aper- Schichten of the Jurassic, near Buren (Canton
ture to be loop-shaped, extending high in the aper- Aargau), Switzerland. In his doctoral dissertation
tural face, sited on the anterior side of the very (1881a), however, no type locality was designated.
small umbilicus, just like the apertures of Oesterle (1968) designated the 'Eisengraben'
Bathonian and Callovian species of Globuligerina. section (80 km NW of Zurich, Canton Aargau) of
However, unlike those species, the surface appears the Birmensdorfer Schichten as the type locality
to lack, in all the previously published and our for all foraminifera species described by Haeusler
new SEM images, any prominent pseudomuricae; from those beds. The topotypes illustrated here
the surface can even appear to be smooth, even (Plate 2.6, Figs 1-8) were collected by Stam
though the specimens are clearly not internal (1986, p. 111) from the Eisengraben section, the
moulds. The shape of the test of G. calloviensis same layer as Oesterle's sample no. 4; 'they
and the characters of its aperture are both very are rare and small in size, [and] this sample
close indeed to G. oxfordiana (Grigelis). Either contained well preserved planktonics'. Earlier,
these two species will prove to be synonyms or Oesterle (1969) had attempted to revise the work
G. calloviensis is the immediate ancestor to the of Haeusler and redescribed 'G.' helvetojurassica
heavily muricate G. oxfordiana (in which case it from the Renggeri Tonen of Liesberg (75 km west
may even be considered to be a subspecies of G. of the Eisengraben section). These were clearly
oxfordiana). We are retaining both taxa at specific pyrite moulds as were Masters' (1977, plate 21,
rank because their stratigraphic ages are different figs 2-4) illustrated SEM images (called
although consecutive; this is open to revision. 'Globigerina? helvetojurassica') from the Oxford-
ian of the Birmensdorfer Beds of Eisengraben. In
Range: Early to early Late Callovian of the contrast, the specimens from the Eisengraben
Crimea, Ukraine. section collected by Starn (1986, plate 7, figs 6--12;
plate 8, figs 1-3) from the Birmensdorfer
Genus Haeuslerina gen.nov. Schichten, Transversarium Zone ('Middle'
Type species Globigerina helvetojurassica Oxfordian) were perfectly preserved. Some of
Haeusler, 1881 Starn's figured specimens have been re-imaged
here (Plate 2.6, figs 1-8). All are covered in scat-
Diagnosis: Conoglobigerinidae with an aperture tered pseudomuricae, not aligned into ridges, and
which is intraumbilical-extraumbilical in extent, very distinctively possess very small umbilici and
reaching almost to the periphery of the penultimate primary apertures which were clearly intra-
whorl, and in which the chamber walls are of extraumbilical.
normal conoglobigerinid thickness and the inter- Starn (1986, pp. 110, 111) did not notice the
cameral sutures are depressed. [Gender: feminine.] difference in the extent of the aperture in his spec-
imens compared to that of Globuligerina oxfor-
Haeuslerina helvetojurassica (Haeusler, 1881) diana (Grigelis), and he mistakenly considered the
(Plate 2.6, Figs 1-8; Figure 2.2) topotypes of 'G.' helvetojurassica to be synony-
mous with G. oxfordiana. As Globigerina helveto-
1881a Globigerina helvetojurassica Haeusler, p. jurassica Haeusler was published in 1881, while
36, p1.2, figs 44, 44a. G. oxfordiana Grigelis was not published until
1986 Globuligerina oxfordiana (Grigelis), Stam 1958, the former name would have been the
(pars), pI. 7, figs 6--12 (not figs 1-5), pI. senior one; Stam endeavoured to overcome this
8, figs 1-3 (not pI. 9, figs 1-5, pI. 14, figs by referring Haeusler's name to the International
3,4, 8-15). Commission of Zoological Nomenclature, claim-
ing that G. helvetojurassica should be regarded as
Remarks: Haeusler (1881b) stated that the type nomen oblitum (a forgotten name, ICZN article
 Concluding comments 29

23b) and not used. However, the case was not Diagnosis: A conoglobigerinid with an intra-
followed up by Starn, and the ICZN did not accept extraumbilical aperture and pseudomuricae which
it (Dr P. Tubbs (ICZN), personal communication). fuse into short disconnected ridges, mostly parallel
Now we know that G. helvetojurassica Haeusler to the growth direction. There are no traces of a
belongs to a genus (Haeuslerina gen.nov.) distinct reticulation forming in the surface sculpture.
from Globuligerina. The specimens figured by [Gender: feminine.]
Masters (1977, pI. 21, figs 2-4) and Oesterle
Compactogerina stellapolaris (Grigelis, 1977)
(1968, text-figs 51, 52a) were internal moulds,
probably like the specimens in Haeusler's own (Plate 2.7, Figs 1-7, Plate 2.8, Fig. 1; Figure
collection (Masters, 1977, p. 459). Therefore, the 2.2)
specimens figured for this chapter, like those
1977 Globuligerina stellapolaris Grigelis (in
figured by Starn (1986, plate 7, figs 6-12, plate 8,
Grigelis et al.), p. 927, text-figs la-c.
figs 1-3) are those which must now be used for
the interpretation of the morphology of
Haeuslerina helvetojurassica (Haeusler). Remarks: The holotype figured by Grigelis
Although Haeusler (1881a; 1881b) did not (1977) is a specimen with a very compact test, in
choose a holotype for his species, the existence of which the sutures are only very slightly depressed
syntypes (referred to by Masters, 1977), albeit and the equatorial periphery is not lobulate. This
internal moulds, unfortunately makes it impossible morphology agrees with that of two metatypes
to designate a neotype (ICZN, article 75). illustrated here (Plate 2.7, Figs 1-3,7). The sutures
of another topotypic specimen also illustrated on
Plate 2.7 (Figs 4-6), however, are much more
Range: This taxon is known only from the
depressed and the equatorial periphery is more
Oxfordian of south central Europe.
lobulate; if the specimens figured here are indeed
Haeuslerina parva (Kuznetsova, 1985) all conspecific, then they give a view of the range
of the variation which occurred in the species, but
(Plate 2.5, Fig. 4; Figure 2.2)
not enough specimens are available to us to be
1985 Globuligerina parva Kuznetsova (in sure. The aperture possesses a narrow lip which
Kuznetsova and Gorbachik), p. 113, pI. 16, partly covers the umbilicus (this lip appears to be
figs 1,2. broken away on Plate 2.7, Fig. 6).

Remarks: This species was poorly illustrated Range: The species was described by Grigelis
when it was first published, the holotype being (1977) from the Early Volgian of the Timan-
shown only by small drawings and a paratype Pechora region in northern Russia (Izhma River,
being imaged by SEM only by its dorsal side. We Pechora River Basin). As such it is the most
illustrate the umbilical side of a metatype, which northerly record of early planktonic foraminifera.
shows the aperture to be intra-extraurnbilical. No The thickened shell and apertural covering
well preserved specimens have yet been found, so may be an adaptation to the relatively cold
the distinction from H. helvetojurassica is not boreal waters of that region, in a manner similar
certain. However, the praemuricae, so conspicuous to modem-day Neogloboquadrina pachyderma
in H. helvetojurassica, appear to be absent in H. (Ehrenberg) .
parva.

Range: Early Kimmeridgian of the Crimea. 2.5 CONCLUDING COMMENTS

Genus Compactogerina gen.nov.
Bullae are present in all of the Favusellacea, from
Type species Globuligerina stellapolaris the earliest to last genera of the Conoglobi-
Grigelis, 1977 gerinidae and Favusellidae, so long as the aper-
30 The Jurassic Favusellacea, the earliest Globigerinina

tures were intraumbilical. When the aperture covering became unnecessary. Bullae are not
became intra-extraumbilical (Haeuslerina, Com- known in the Praehedbergellidae, but umbilical
pactogerina in the Conoglobigerinidae; Ascoliella coverings were developed in the macroperforate
in the Favusellidae) the bullae disappeared, just as descendants of the Hedbergellidae (e.g.
in the Cenozoic Globigerinacea (e.g. Catapsydrax, Rotalipora, Globotruncana, etc.).
with a bulla covering its intraumbilical aperture, The pseudomuricae of the Conoglobigerinidae
compared to Paragloborotalia which has an intra- often fuse to form discontinuous short ridges, and
extraumbilical aperture and no bulla). It may be these are particularly well developed in the strati-
that the bulla was a skeletal structure which graphically youngest species of Conoglobigerina
covered the umbilicus to conceal a mass of (the Berriasian-Early Valanginian species, C.
extrathalamous but digestive cytoplasm analogous caucasica and C. gulekhensis). Lateral fusion of
to that seen today in some living benthic rotalines these ridges form the favose reticulate sculpture
(Alexander and Banner, 1984). It is possible that of Favusella (which first appeared also in the
with the extraumbilical extension of the aperture Berriasian).
and narrowing of the umbilicus this bullate
Plate 2.1
The gamonts of some species of the (normally benthonic) Discorbidae and Cymbaloporidae have a planktonic stage just before
and during reproduction. They independently evolved floatation structures in subglobular chambers at the last stage of growth
of the test.

Figures 1,6 Neoconorbina (Tretomphaloides) concinna (Brady, 1884). Recent, coast of Mombasa, Kenya, nos ZF 4383, ZF
4385: 1, dorsal view (xI65); 6, specimen with outer balloon removed, leaving the imperforate inner float chamber
with a broad tube linking to the spiral whorl (x585).
Figures 2, 3, 7 Cymbaloporetta cifeWi Banner, Pereira and Desai, 1985. Recent, off Mombasa, Kenya, paratypes, no. ZF 4388:
2, dorsal view with chambers alternating in annual series (x75); 3, peripheral view (x85); 7, ventral view
showing uniformly perforate balloon chamber and rimmed apertures (x70).
Figures 4, 8, 9 Cymbaloporetta (Millettiana) millettii (Heron-Allen and Earland, 1915). Recent, off Mombasa, Kenya, nos ZF
4375, ZF 4377, ZF 4378: 4, ventral view with balloon chamber removed, with float chamber plate partly covering
the umbilicus (xI50); 8, oblique ventral view, showing stellate clusters of apertures in the balloon chamber,
arising from the lateral tubes (x150); 9, side view with float chamber broken (x260).
Figure 5 Cymbaloporetta sp. Recent, off Mombasa, Kenya, no. ZF 4379: showing lateral invaginations of float chamber,
some constricted to form vertical tubes (xllO).
All specimens deposited in The Natural History Museum, London (ZF numbers).
Plate 2.2

Figures 1-3,5 Praegubkinella racemosa Wernli, 1995. Early Toarcian, Falciferum Zone, sample W1671, Creux de 1'Ours ,
Teysachaux, Swiss Alps (from Wernli, 1995, pI. 1, figs 1-3, 5), deposited in the Department of Geology and
Palaeontology, Museum d'Histoire Naturelle de Geneve: 1-3, peripheral views; 5, dorsal view (1 is the paratype,
x240); (3 is the holotype, x260).
Figures 6, 8 Praegubkinella fuchsi Wernli, 1995. Early Toarcian, Falciferum Zone, sample W1671, Creux de l'Ours,
Teysachaux, Swiss Alps (from Wernli, 1995, pI. 1, figs 6, 8), deposited in the Department of Geology and
Palaeontology, Museum d'Histoire Naturelle de Geneve: peripheral views (6 is the holotype, x250).
Figures 4, 7, 9 Oberhauserella quadilobata Fuchs, 1967. Early Toarcian, Falciferum Zone, sample W1671, Creux de l'Ours,
Teysachaux, Swiss Alps (from Wernli, 1995, pI. 1, figs 19,24) deposited in the Department of Geology and
Palaeontology, Museum d'Histoire Naturelle de Geneve: 4,9, peripheral views (x250); 7, dorsal view (x250).
Plate 2.3

Figures 1-13 Conoglobigerina dagestanica Morozova, 1961. 1-4, (= Globuligerina arabi Kasimova, 1984). Late Bajocian,
Jurassic, Negramskoye gorge, Aras River, Azerbaijan, deposited in the Azerbaijan Academy of Sciences. Metatype
of G. araksi (specimen diameter 130 /Lm): 1-3, dorsal, peripheral and ventral views (x310); 4, enlargement of the
wall surface to show pseudomuricae often fusing laterally into short ridges (x1500). 5-13, holotype and three
paratypes of C. dagestanica Morozova, 1961, Bathonian, near the villages of Chokh and Gunib, central Dagestan,
deposited in the Collections of the Geological Institute of the Academy of Sciences of Russia, Moscow - holo-
type (no. 3513/2) (specimen diameter 190 /Lm): 5-7, dorsal, peripheral and ventral views (x21O); paratype
(specimen diameter 150/Lm): 8-10, dorsal, peripheral and ventral views (x265); paratype (specimen diameter
190 /Lm): 11, 12, enlargement of the surface showing microperforations and pseudomuricae (xI250) and dorsal
view (x210); paratype (no. 3513) (specimen diameter 210 /Lm): 13, peripheral view (xI9O).
Plate 2.4

Figures 1-4, 6 Conoglobigerina avarica Morozova, 1961. Early Bathonian, central Dagestan, holotype and three paratypes
deposited in the Collections of the Geological Institute of the Academy of Sciences of Russia, Moscow -
holotype (no. 3513/3) (specimen diameter (height of coil) 170 fLm): I, peripheral view showing a very high
spire (x235); paratype (specimen diameter (height of coil) ISO fLm): 2,3, ventral (x330) and peripheral views
(x265); paratype (no. 3513/24): 4, enlargement of the surface (x 1100); paratype (no. 3513/19) (specimen
diameter 180 fLm): 6, dorsal view (x220).
Figures 7-12, 16 Conoglobigerina balakhmatovae (Morozova, 1961). Early Bathonian, central Dagestan, holotype and three
paratypes deposited in the Collections of the Geological Institute of the Academy of Sciences of Russia,
Moscow - paratype (no. 3514): 7, enlargement of the surface showing microperforations and pseudomuricae
(xI500); paratype (no. 3513/9) (specimen diameter 120 fLm): 8, 12, peripheral view (showing a low spired
test with a slightly convex dorsal side) and ventral views (x335); holotype (no. 3513/1) (specimen diameter
140 fLm): 9-11, dorsal, peripheral and ventral views (x285); paratype (specimen diameter 180 fLm): 16, dorsal
view showing a fast rate of chamber enlargement (x220).
Figures 5, 13-15 Conoglobigerina dagestanica Morozova, 1961. 5 (= C. avarica Morozova, 1961 (pars., Early Bathonian,
central Dagestan, deposited in the Collections of the Geological Institute of the Academy of Sciences of
Russia, Moscow; paratype of C. avarica (specimen diameter 140 fLm): ventral view (x285). 13-15, (=
Globigerina (Eoglobigerina) balakhmatovae Morozova, 1961 (pars.), pI. I, figs 9-11, paratype only),
Bathonian, the village of Chokh, central Dagestan, deposited in the Collections of the Geological Institute
of the Academy of Sciences of Russia, Moscow; same paratype (specimen diameter 150 fLm): dorsal, periph-
eral and ventral views showing four chambers in the final whorl (x265).
Plate 2.5

Figures 1-3 Conoglobigerina meganomica (Kuznetsova, 1980). Late Callovian, northern Crimea, deposited in the Collections
of the Geological Institute of the Academy of Sciences of Russia, Moscow, paratype (specimen diameter
210 11m): dorsal, peripheral and ventral views (xI90).
Figure 4 Haeuslerina parva (Kuznetsova, 1985). Early Kimmeridgian, Crimea, deposited in the Collections of the
Geological Institute of the Academy of Sciences of Russia, Moscow, metatype (specimen diameter 140 11m):
ventral view showing an intra-extraumbilical aperture (x285).
Figures 5-10 Globuligerina calloviensis Kuznetsova, 1980. Early-Middle Callovian of the Crimea, Ukraine, two metatypes
deposited in the Collections of the Geological Institute of the Academy of Sciences of Russia, Moscow, metatype
(specimen diameter 110 11m): 5-7, dorsal, peripheral views, and ventral view showing the aperture to be
Globuligerina-like (loop-shaped) (x365); metatype (specimen diameter 180 11m): 8-10, dorsal, peripheral and
ventral views (x220).
Figures 11-14 Favusella hoterivica (Subbotina, 1953). Hauterivian, northern Caucasus, deposited in the All-Union Petroleum
Scientific Research Prospecting Institute (VNIGRI), St Petersburg, Russia, holotype (no. 5165) (specimen diam-
eter 145 11m): enlargement of the wall surface showing fine reticulation (xllOO), dorsal, peripheral and ventral
views (x275).
Plate 2.6

Figures 1-8 Haeuslerina helvetojurassica (Haeusler, 1881). Oxfordian, Eisengraben section of the Birrnensdorfer Schichten,
Canton Aargau, Switzerland, three topotypes collected and figured by Stam (1986), deposited in Atlantic
Geoscience Centre, Bedford, Canada; topotype no. GSC 113076 (specimen diameter 110 !Lm): 1-3, dorsal,
peripheral and ventral views showing an intra-extraumbilical aperture (x365), re-imaged, previously figured by
Stam, 1986, pI. 7, fig. 12); topotype no. GSC 113077 (specimen diameter 100 !Lm): 4--6, enlargement of surface
showing scattered pseudomuricae (x2000), dorsal and peripheral views (x400), re-imaged, previously figured
by Stam, 1986, pI. 8, figs 1-3); topotype no. GSC 113074 (specimen diameter 100 !Lm): 7,8, peripheral (x400)
and oblique-ventral (x450) views, re-imaged, previously figured by Starn, 1986, pI. 7, figs 6,7).
Figures 9-12 Conoglobigerina caucasica (Gorbachik and Poroshina, 1979). Early Berriasian, village of Ugakh, SE Caucasus,
Azerbaijan, two metatypes deposited in the Geological Faculty of Moscow State University, Russia - metatype
(specimen diameter 160 !Lm): 9-11, dorsal view with a low spire, peripheral view showing a convex dorsal
side, and ventral view with an intraumbilical aperture which unfortunately here has a sediment covering its
umbilicus (x250); metatype: 12 enlargement of the wall surface (dorsal side) showing pseudomuricae some-
times fusing into discontinuous, blunt ridges (x1100).
Figures 13-15 Conoglobigerina gulekhensis (Gorbachik and Poroshina, 1979). Early Berriasian, village of Gulekh, SE Caucasus,
Azerbaijan, deposited in the Geological Faculty of Moscow State University, Russia, metatype (specimen diam-
eter 170 !Lm): dorsal view, peripheral view showing a convex dorsal side, and ventral view with intraumbilical
aperture (x235).
Plate 2.7

Figures 1-7 Compactogerina stellapolaris (Grigelis, 1977). Early Vo1gian, exposure on the left bank of the Izhma River,
near the village of Zagrivochnaya, Pechora River Basin, northern Russia, three metatypes deposited in the
Geological Faculty, Moscow State University, Russia - metatype (specimen diameter 235 fJ-m): 1-3, dorsal,
peripheral and ventral views (x170); metatype (specimen diameter 265 fJ-m): 4-6, dorsal, peripheral and ventral
views (x150); metatype (specimen diameter 260 fJ-m): 7, ventral view (xI55).
Figures 8-10 Conoglobigerina avariformis Kasimova, 1984. Late Bajocian, Negramskoye gorge, Aras River, Nakhichevan,
Azerbaijan, donated by Dr G.K. Kasimova to The Natural History Museum, London, metatype (specimen
diameter 320 fJ-m): dorsal, peripheral and ventral views (xI25).
Figures 11-14 Conoglobigerina jurassica (Hofman, 1958). Bathonian-Early Callovian, southeastern Crimea, deposited in the
Geological Faculty, Moscow State University, Russia; neotype, designated herein (specimen diameter 270 fJ-m):
11-13, dorsal, peripheral and ventral views (xI50); 'paratype' (specimen diameter 260 ""m): 14, dorsal view
(xI55).
Plate 2.8

Figure I Compactogerina stellapolaris (Grigelis, 1977), Early Volgian, Izhma River, Pechora River Basin, northern Russia,
deposited in the Geological Faculty, Moscow State University, Russia, metatype: enlargement of the wall surface
showing microperforations and pseudomuricae (x650),
Figure 2 Conoglobigerina avariformis Kasimova, 1984, Bajocian, Negramskoye gorge, Aras River, Nakhichevan,
Azerbaijan, donated by Dr G.K. Kasimova to The Natural History Museum, London, metatype: enlargement of
the wall surface showing pseudomuricae (x750).
Figure 3 Conoglobigerina jurassica (Hofman, 1958). Bathonian-Early Callovian, SE Crimea, deposited in the Geological
Faculty, Moscow State University, Russia, 'paratype': enlargement of the surface showing pseudomuricae (x850).
Figures 4-12 Globuligerina bathoniana (Pazdrowa, 1969). Middle Bathonian, Ogrodzieniec, Poland, two metatypes deposited
in the Geological Faculty, Moscow State University, Russia - metatype (specimen diameter 160 [Lm): 4-6, dorsal,
peripheral and ventral views (x250), 10, II, same specimen, two views of surface detail showing pseudomuricae
usually separate but some adjacent pairs fusing into very short and discontinuous ridges (x1250 and xl 100, respec-
tively). Same locality, metatype donated by Dr B. Masters to The Natural History Museum, London (specimen
diameter 170 [Lm): 7-9, dorsal, peripheral and ventral views (x235); topotype: 12, enlargement of the apertural
face showing a low loop-shaped aperture (x750). Note, traces of a bulla can be seen on Figs 6 and 9.
Plate 2.9

Figures 1-15 Globuligerina oxfordiana (Grigelis. 1958). Oxfordian, Makarievo, Upper Volga Basin, Russia, deposited in the
Geological Faculty of Moscow State University, Russia (specimen diameter 150 f,Lm): 1-3, dorsal, peripheral and
ventral views, the trace of the bulla is very clear (x265). Oxfordian, near Moscow, Russia, deposited in the
Geological Faculty of Moscow State University (specimen diameter 140 f,Lm): 4, ventral view (x285). Oxfordian,
near Le Havre, N France (donated to The Natural History Museum, London, by G. Bignot) (specimen diameter
130 f,Lm): 5-7, 9, 10, dorsal, peripheral (x285) and oblique-ventral (x330) views, detail of surface (xI5OO) and
loop-shaped aperture (x1250), respectively; 11, another specimen, detail of surface showing some pseudomuricae
fusing into short ridges (xIOOO). Oxfordian, near Moscow, Russia, deposited in the Geological Faculty of Moscow
State University (specimen diameter 130 f,Lm): 8, ventral view showing a bulla (x300). 12-15, Oxfordian,
Makarievo, Upper Volga Basin, Russia, donated by T. Gorbachik to The Natural History Museum, London, nos
PF62915, PF 62916: 12-14, dorsal, peripheral and ventral views (x235; specimen diameter 170 f,Lm); 15, ventral
view (x285, specimen diameter 140 f,Lm).
Plate 2.10

All Cenozoic contaminants!

Figures 1-7 Globigerina spuriensis Bars and Ohm, 1968. Said to be Late Bajocian-Early Bathonian but believed by us to
be Cenozoic (Palaeogene) contaminants, Rocchetta section, Noce River, N Italy, deposited in the Senckenberg
Museum, Frankfurt, Germany, no. SMF XXVII 5830, paratype: 1-3, dorsal, peripheral and ventral views (x90;
specimen diameter 440 /Lffi); no. SMF XXVII 5831, holotype: 4-7, dorsal, peripheral and ventral views (x70) ,
and detail of surface showing deep macroperforations (x550) of a typical Cenozoic form (?Catapsydrax), spec-
imen diameter 570 110m.
Figures 8-11 Globigerina gaurdakensis Balakhmatova and Morozova, 1961. Claimed to be Late Bajocian, from the Daurdak
region of Turkmenia, but believed by to us to a contaminant, deposited in the FN. Chemyshev Museum, A11-
Union Scientific Research Geological Institute, St Petersburg, Russia, holotype (no. 9359/2) (specimen diam-
eter 155 110m); dorsal, peripheral and ventral views (x260), and enlargement of wall surface showing the typical
deep macroperforations (xl100) of a Cenozoic species.
Figures 12-14 Globuligerina geczyi Goriig, 1994. Said to be Early Jurassic (Hettangian-Sinemurian) but believed by us to be
Palaeocene in age, from the Gerecse Mountains, N Hungary (from Goriig, 1994, pI. 1, figs 1, 6, 9), deposited
in the Hungarian Natural History Museum, Budapest, paratype M.93.52: 12, dorsal view (xI95; specimen
diameter 205 110m); holotype, M.93.51: 13, peripheral view (xI75; specimen diameter 230 110m); paratype,
M.93.50: 14, ventral view showing muricae spread evenly over the test (xI95; specimen diameter 205 110m). We
believe figs 12, 14 to belong to a Praemurica, and fig. 13 to be of a Globoconusa daubjergensis (Briinnimann,
1953). (From Gorog, 1994, pI. 21, figs 1,6,9).
 3
The Early Cretaceous
families
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

3.1 INTRODUCTION Aptian. Gorbachikella seems to have evolved from

Conoglobigerina in the Late Valanginian.
All Jurassic planktonic foraminifera belong to the Maamouri and Salaj (1995) described five new
Favusellacea, a muricate, microperforate and prob- species (one of them belonging to a new genus)
ably aragonitic group. In the Early Cretaceous, from the upper Valanginian deposits of Jebel Oust
the first true Globigerinacea developed with the in Tunisia. These forms are the first ancestors of
Praehedbergellidae and Schackoinidae, which Hauterivian Gorbachikella. Maamouri and Salaj
were microperforate, non-muricate, non-pustulate (1995) referred to them as Globuligerina neili, G.
and non-spinose and calcitic families. The test spiralis, G. mejezensis, G. compressa and
surface of the Praehedbergellidae is either smooth Archaeokassabella biapertura. The specimens
or furnished with perforation cones (unlike any figured by Maamouri and Salaj are badly
other Globigerinacea but reminiscent of some preserved and dirty, as our Gorbachikella speci-
Heterohelicacea, such as Cassigerinella, Guembe- mens used to be (Plate 6.1, Figs 5, 6) before we
litria, etc.). These eventually gave rise to the devised the new method of cleaning the separated
Hedbergellidae and Planomalinidae (macroper- and picked specimens (which consisted of washing
forate and muricate) and the different lineages of them individually in ultrasound with a sodium
the Late Cretaceous planktonic foraminifera. The hexametaphosphate [NaP03)nNazOl solution, and
Praehedbergellidae and Schackoinidae typify the with very successful results, as shown in Plate 6.1,
holoplanktonic foraminiferal fauna of the Early Figs 3, 4. None of the specimens figured by
Cretaceous, and were the first geographically Maamouri and Salaj is muricate, or have the bullae
widespread Globigerinacea (Banner and Desai, characteristic of the Jurassic Globuligerina and the
1988). These families are known throughout the Cretaceous Favusella. They must be smooth. They
tropical-subtropical Early Cretaceous world. are either the youngest known Conoglobigerina or
The stratigraphically earliest genus of the Prae- the oldest Gorbachikella. They may be interme-
hedbergellidae was Gorbachikella, known from diate forms, showing the evolution from
the Late Valanginian to the end of the Early Conoglobigerina to Gorbachikella; however, we
54 The Early Cretaceous families

regard them as belonging to the oldest known Gorbachikella as it evolved from the ancestor
species of Gorbachikella as they show no distinc- Conoglobigerina. We have seen deformed speci-
tive morphocharacters to separate them from this mens from the Early Barremian of Tunisia, but
genus. The types of 'Globuligerina' neW figured they are not known to have an additional primary
by Maamouri and Salaj (1995, plate 1, figs 1-5) aperture on the spiral side (Plate 6.2, Figs 7-9).
have a low aperture bordered with a lip, small
umbilicus, depressed chambers and are considered
here as Gorbachikella neW (Plate 6.6, Figs 7, 8). 3.2 KEY TO THE SUPRAGENERIC TAXA
It ranges from the Late Valanginian to the end of
the Early Barremian. The figures of Order GLOBIGERININA Delage and Herouard,
'Globuligerina' spiralis are very dirty and the 1986
early whorls are not discernible in the holotype;
in the paratype (Maamouri and Salaj, 1995, plate Superfamily FAVUSELLACEA Longoria, 1974
4, fig. 3) there are five chambers in the first whorl. (nom.corr.) emend. Banner and Desai, 1988
The only known Hauterivian Gorbachikella with
five chambers in the first whorl is Gorbachikella Family FAVUSELLIDAE Longoria, 1974
grandiapertura BouDagher-Fadel et al., 1995,
Test trochospiral with two or more whorls;
which also has a high spire and depressed cham-
initially the first whorl with four or more cham-
bers when viewed from the spiral side, like those
bers, final whorl of about four chambers, rarely as
of the paratypes of G. spiralis. The originally
many as seven; spiral side evolute, umbilical side
figured specimens of G. spiralis have a primary
involute with a small umbilicus; aperture intra-
aperture as high as the typical Gorbachikella
umbilical, rarely intra-extraumbilical, a high or
grandiapertura. We suggest, therefore, that G.
low arch, which may be strongly asymmetric; wall
spiralis should be referred to as Gorbachikella
microperforate (external diameters <1.5 !-Lm),
grandiapertura, the oldest known Gorbachikella.
covered with rounded pseudomuricae which fuse
There are eight figures of G. mejezensis
into ridges which in tum anastomose to form retic-
(Maamouri and Salaj, 1995, plate 3, figs 1,2; plate
ulations over the test surface; wall is believed to
5, figs 1--6). They are again very dirty and none
be composed primarily of aragonite.
of them shows the umbilical side and the aperture
of this species. The spiral side is as high as that
Key to the genera of the FAVUSELLIDAE
of Gorbachikella depressa BouDagher-Fadel et
al., 1995, and we suggest that it should also be Four to five chambers in the final whorl; aperture
referred to as the latter (which has priority) a low or moderately high symmetrical arch,
(Chapter 6) until we can examine (and maybe usually with a narrow lip, sometimes with a bulla-
clean) more specimens. On the other hand, the like last chamber covering the umbilicus; wall
figures of 'Archaeokassabella biapertura', from microperforate; surface covered by anastomosing
the Upper Valanginian deposits of Jebel Oust ridges which enclose groups of microperforations
(Plate 6.2, Figs 10, 11), are not good and there is in a favose reticulation; pseudomuricae may be
no clear spiral or umbilical view, but the genus is present at the confluences of the anastomosing
supposed, by its authors, to possess an additional ridges. The continuous reticulation of the surface
(supplementary, sutural) aperture on the spiral ridges distinguishes Favusella from Conoglobi-
side. Four figures are oblique. Only five specimens gerina and Globuligerina. Earlier species of
were found by Maamouri and Salaj. We suspect Favusella (e.g. hoterivica) have pseudomuricae
that these are distorted and deformed specimens fused into low ridges which weakly anastomose
of an early Gorbachikella. However, if they are forming a low reticulation of weak: rounded ridges
proven to be normal representatives of a new and narrow concave depressions between them.
taxon, Archaeokassabella biapertura would be On the other hand, the youngest Favusella have
a very early and short lived off-shoot from narrow ridges and broad, flat depressions.
 Key to the suprageneric taxa 55

1 Aperture intraumbilical; ~~ chambers in the Genus Gorbachikella

last whorl; Berriasian-Early Cenomanian. Banner and Desai, 1988
Genus Favusella Michael, 1973 (Type species Gorbachikella kugleri Bolli, 1959)
(Type species Globigerina washitensis 1.1' Smooth but sometimes with perforation
Carsey, 1926) cones, aperture umbilical-extraumbilical:
2 Aperture intra-extraumbilical; 5-7 chambers in 1.1'.1 ~~ chambers per whorl:
the final whorl; Late Aptian-latest Albian. 1.1' .1.1 Chambers of last whorl glob-
Genus Ascoliella Banner and Desai, 1988 ular or slightly depressed; Hauterivian-
(Type species Ascoliella scotiensis Late Aptian
Banner and Desai, 1988) Genus Praehedbergella Gorbachik
and Moullade, 1973
Superfamily GLOBIGERINACEA Carpenter, (Type species Globigerina tuschepsensis
Parker and Jones, 1862 Antonova, 1964)
Test trochospiral, or planispiral, calcitic 1.1' .1.2 Chambers of last whorl elon-
gate, higher than long; Barremian
1 Microperforate, lacking muricae, pseudo- Genus Lilliputianelloides gen. nov.
muricae, pustules or spine-bases, non-pustulate, (Type species Clavihedbergella
nonspinose, but sometimes possessing perfora- eacretacea Neagu, 1975)
tion cones especially near the aperture 1.1'.2 5 or more chambers per whorl:
1.1 Test trochospiral, primary aperture is intra- 1.1' .2.1 Chambers subglobular to de-
umbilical or intra-extraumbilical pressed; Barremian-Early Danian
Family PRAEHEDBERGELLIDAE Genus Blefuscuiana Banner
Banner and Desai, 1988 and Desai, 1988
1.2 Test planispiral, biumbilicate; aperture (Type species Blefusciana kuznetsovae
interiomarginal, equatorial, symmetrical, Banner and Desai, 1988)
extending laterally into each umbilicus 1.1'.2.2 Chambers radially elongate;
Family SCHACKOINIDAE Pokorny, 1958 latest Barremian-Late Aptian
2 Test trochospiral, macroperforate (as opposed Genus Lilliputianella Banner
to microperforate), muricate, aperture intra- and Desai, 1988
extraumbilical (Type species Lilliputianella langorii
Family HEDBERGELLIDAE Banner and Desai, 1988)
Loeblich and Tappan, 1961 2 Initially trochospiral, later whorls streptospiral;
3 Test planispiral, macroperforate, muricate; Late Aptian
aperture interiomarginal, equatorial, symmet- Genus Wondersella Banner and Strank, 1987
rical, extending laterally into each umbilicus, (Type species Wondersella athersuchi
final aperture and relict apertures are usually Banner and Strank, 1987)
furnished with portici
Family PLANOMALINIDAE
Loeblich and Tapan, 1957, emended Key to the genera of the SCHACKOINIDAE
1 4-5 chambers per whorl; chambers longer than
Key to the genera of the or as long as high, not radially and narrowly
PRAEHEDBERGELLIDAE elongate; portici usually broadest at their poste-
rior or at their extreme anterior; latest Early
1 Trochospiral throughout Barremian-Late Aptian
1.1 Smooth, aperture intraumbilical; 4 (rarely Genus Blowiella Kretzchmar and Gorbachik,
5 chambers) in the first whorl, 4 chambers 1971, emend. Banner and Desai, 1988
in the final whorl; Late Valanginian-Early (Type species Planomalina blowi Bolli, 1959)
Aptian 2 ~~ chambers per whorl; chambers radially
56 The Early Cretaceous families

elongate, higher than long, but broadly clavate; Bronnimann and Brown, 1958 (Late Aptian-top
latest Early Aptian-Late Aptian (? Early Aptian) of Maastrichtian) evolves from Blefuscuiana by
Genus Claviblowiella BouDagher-Fadel acquiring macroperforations (as opposed to
et al., 1996 microperforations), and later muricae. The muricae
(Type species Claviblowiella of Hedbergella fuse even later in the Albian
saundersi Bolli, 1975) in order to form the peripheral keel of
3 4-6 chambers per whorl; with later chambers Praeglobotruncana (Albian-Early Turonian), a
becoming radially and broadly elongate and ter- morphocharacter unknown in the Praehed-
minally bulbiform; Late Barremian-Early Albian bergellidae.
Genus Leupoldina Bolli, 1957, emend.
Banner and Desai, 1988
Key to the genera of the
(Type species Leupoldina protuberans
PLANOMALINIDAE
Bolli, 1957)
4 4-6 elongated chambers in the last whorl, each 1 Test nearly involute; macroperforate, some-
ending with one narrow tubulospine, occasion- times the macroperforations become sunk into
ally two, at the end of last chamber, Late Aptian perforation depressions (Plate 12.1, Fig. 3),
Maastrichtian sometimes the wall is muricate in early whorls;
Genus Schackoina Thalmann, 1932 periphery rounded, sometimes compressed in
(Type species Siderolina cenomana the early stage but never developing a keel;
Schacko, 1897) Late Aptian-Early Cenomanian
5 8-10 chambers in the last whorl, broad umbilici Genus Alanlordella
and longer portici; chambers compressed latter- BouDagher-Fadel, 1996
ally but not radially elongate; Late Aptian- (Type species Alanlordella banneri
Early Albian BouDagher-Fadel, 1996)
Genus Globigerinelloides Cushman and 2 Test nearly involute; macroperforate, muricate,
Ten Dam, 1948 the muricae fusing at the periphery and on the
(Type species Globigerinelloides algerianus sutures to form a murico-carina; periphery
Cushman and Ten Dam, 1948) compressed; latest Albian-Late Cenomanian
Genus Planomalina Loeblich and Tappan,
Family HEDBERGELLIDAE Loeblich and
1946, emended
Tappan, 1961
(Type species Planomalina buxtorji
In the earliest Late Aptian Hedbergella Gandolfi, 1942)
 4
The Favusellidae - the
Cretaceous acme of
the Favusellacea
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

4.1 INTRODUCTION The cells within the reticulations of Favusella

were initially small and their bases were curved
In the Early Berriasian to Early Valanginian of and concave. As the genus evolved, the reticula-
southeastern Europe, species of Conoglobigerina tions became broader and flatter, culminating in
occur which are covered in pseudomuricae. These the form displayed by F. washitensis (Carsey) in
pseudomuricae often fused together to form short, the Late Aptian-Early Cenomanian. Therefore, the
discontinuous ridges. The species were the high gradation from finely reticulate Berriasian-
spired C. gulekhensis and the lower spired C. Hauterivian forms of Favusella to broadly reticu-
caucasica (both described by Gorbachik and late forms in the Albian-Early Cenomanian
Poroshina, 1979), and these may constitute the last appears to be continuous and complete.
dimorphic pair of the essentially Jurassic genus, All species of Favusella possessed four (occa-
Conoglobigerina. sionally three and a half) chambers visible on the
In the Berriasian, these discontinuous ridges umbilical side of the last whorl and an intra-
fused to form a reticulate, pseudomuricate coating umbilical aperture. Occasionally, the aperture
over the surface of the test. This happened in the was covered by a bulla (as in the neotype of
oldest species which we would refer to the genus F. washitensis proposed by Longoria, 1974);
Favusella (Figure 4.1). No dimorphic pairs are similar bullae occurred in Globuligerina and
known in this new genus and its species became Conoglobigerina, and differ from the bullae of
worldwide. Consequently, we believe that a holo- Cenozoic species in often possessing short discon-
planktonic mode of life had at last been achieved tinuous ridges formed from pseudomuricae. All
in this evolutionary lineage (although the separate these bullae allowed contact with the exterior only
and stratigraphically earlier Globuligerina had through a single accessory, infralaminal aperture
also become holoplanktonic). at their margins; the bullae of Catapsydrax in the
 Co)
...
Co) ~ ~
... <0
~ ...., !:
......., .......,
~ m
......
N

BERRIASIAN VALANGINIAN HAUTERIVIAN BARREMIAN APTIAN ALBIAN CENOMANIAN

EARLY LATE EARLY LATE EARLY LATE EARLY LATE ~ARLY LATE EARLY LATE EARLY LATE
~.
Favusella h/ltermann/

F. hater/vlea

F. st/ft/a

F. wash/tens/s

Ascallella n/t/da

A.quadrata

A. sc/tu/a

A. va/ash/nae

_ _ . ____ L - ~.

- - - ------- - - - -

Figure 4.1 Stratigraphical distribution of the Favusellidae (Favusellacea) - Favusella and Ascoliella. Chronostratigraphy and Absolute Time Scale after Gradstein et al. (1995).
 Systematics 59

Cenozoic were perforate, not muricate and had Type species Globigerina washitensis Carsey,
from one to four accessory, infralaminal apertures. 1926
Favusella orbiculata Michael, 1973, p. 214,
Favusella washitensis (Carsey, 1926)
plate 4, figs 1-3 (Plate 4.3, figs 4--6) and F.
wenoensis Michael, 1973, p. 216, plate 5, figs 4-9; (Plate 1.2, Fig. 6, Plate 4.3, Figs 7-10, Plate
plate 7, figs 4, 5 (Plate 4.6, figs 4-7) are both 4.4, Figs 1-9; Figure 4.1)
nomina dubia, deformed specimens, and we do not
consider them valid. The type figure of Favusella 1926 Globigerina washitensis Carsey, p. 44, pI.
planata Longoria and Gamper, 1977, pp. 207, 208 7, fig. 10.
was not given and the type level was not desig- 1931 Globigerina washitensis Carsey; Plummer,
nated, except that it occurred in the Late Albian p. 193, pI. 13, figs 12a, b (first neotype).
of the Sabinas Basin of northern Mexico. How- 1961 Hedbergella washitensis (Carsey);
ever, the type description of F. planata (six to Loeb1ich and Tappan, pI. 4, fig. 11.
seven chambers in the last whorl and an intra- 1963 Reticuloglobigerina Reiss, p. 74.
extraumbilical aperture) seems to place this 1973 Favusella washitensis (Carsey), Michael,
species firmly in the Ascoliella. pp. 212, 213, pI. 5, figs 1-3.
In the Late Aptian, the genus Ascoliella evolved 1974 Favusella washitensis (Carsey), Longoria,
from Favusella by extension of the aperture pp. 74, 75, pI. 26, figs 4-6 (second neotype).
towards the periphery. This intra-extraumbilical 1974 Favusella sp. Longoria, p. 74, pI. 26, figs
aperture allowed the species of Ascoliella to 8, 9.
acquire five or six chambers in the last whorl, but 1977 Favusella confusa Longoria and Gamper,
no bullae are known to have occurred. In the pp. 204, 207, pI. 3, figs 4--6 (paratype),
Conoglobigerinidae, intra-extraumbilical apertures 10-12; pI. 4, figs 25-27 (holotype).
were formed in the genera Haeuslerina and 1977 Globigerina washitensis Carsey; Masters,
Compactogerina, but these taxa are not known to p. 477, pI. 25, fig. 4; pI. 26, figs 1-3 (topo-
have acquired the large number of chambers in type).
the final whorl displayed by species of Ascoliella. 1982 Reticuloglobigerina washitensis (Carsey),
Koutsoukos et al. (1989) suggested that all the Banner, p. 188, fig. 5.5.
various Favusella taxa were ecophenotypic 1985 Favusella washitensis (Carsey), Caron,
expression of a single stock suited to shallow, p. 45, Figs 25.25, 26.
warm, hypersaline, carbonate-saturated environ- 1988 Favusella washitensis (Carsey), Banner
ments. We do not agree, because the taxa and Desai, pp. 148-150, pI. 2, figs 6,7.
described below are known to have different strati-
graphic ranges and different regional distributions Remarks: F. washitensis has commonly four
(Ascoliella, for example, is not known in Tethys chambers in the last whorl, but occasionally only
even though the stratigraphically latest species of three and a half chambers are visible on the umbil-
Favusella is geographically widespread). ical side. An intraumbilical aperture occurs with
a microperforate wall which has the surface
covered by anastomosing ridges, which enclose
4.2 SYSTEMATICS groups of the microperforations in the cells of the
favose reticulation. Pseudomuricae mayor may
Superfamily FAVUSELLACEA Longoria, 1974 not be present at the confluences of the anasto-
(nom.corr.) emend. Banner and Desai, 1988 mosing ridges; Plate 4.4, figs 8-9 shows how
incipient pseudomuricae are preserved at the inter-
Family FAVUSELLIDAE Longoria, 1974 section of the ridges even in the stratigraphically
youngest form of the evolutionary lineage,
Genus Favusella Michael, 1973 recalling its conoglobigerine ancestor.
60 The Favusellidae

The type specimens of F. washitensis, from the Favusella hiltermanni (Loeblich and Tappan,
Washita group, Texas, were lost from the Univer- 1961)
sity of Texas Micropalaeontological Collection and (Plate 4.7, Figs 4-6; Figure 4.1)
a neotype was proposed by Plummer (1931) from
1961 Hedbergella hiltermanni Loeblich and
the Del Rio Formation (Early Cenomanian), of the
Tappan, p. 275, pI. 4, figs 13a-c (?12a-c).
Washita Group, at Shoal Creek, Austin, Texas.
Longoria (1974) noted that Carsey's original Remarks: This species has five chambers in the
photograph was a dorsal view, showing four and a final whorl visible dorsally, while four abut
half chambers dorsally in the last whorl of the only the umbilicus on the ventral side; the umbilicus is
figured syntype. Consequently, Longoria (1974) deep and narrow, and is sometimes covered by
believed it was significantly different from the neo- a bulla, as in the holotype. The surface of the
type figured by Plummer (1931), as the latter chamber is coarsely reticulate, and the bulla has
possessed only four chambers visible dorsally in a weaker reticulation.
the last whorl. Therefore, Longoria proposed a new The reticulate favose pattern of F. hiltermanni
neotype from the Grayson Formation of the same is much coarser than that of F. washitensis
locality, which had four and half chambers visible (Carsey) and the cells are broader and their floors
dorsally in the last whorl. The specimens with only are very flat; the umbilicus of the holotype of
four chambers in the last whorl (seen dorsally) were F. hiltermanni is very narrow.
named by Longoria and Gamper (1977) as the new
species F. confusa. It is interesting to note that the Range: Early Cenomanian. This species was
specimen figured by Tappan (1940), from the Late first described from the Zeltberg Brickworks, at
Albian Grayson Formation of north Texas, had four Luneberg, southeast of Hamburg, Germany.
chambers visible dorsally in the last whorl, while
Favusella hoterivica (Subbotina, 1953)
that figured by Loeblich and Tappan (1961), a topo-
type from the Cenomanian of the Del Rio Clay, (Plate 2.5, Figs 11-14; Plate 4.1, Figs 1-17;
Austin, Texas had four and a half chambers. It is Plate 4.2, Figs 1-8; Figure 4.1)
clear from these and other topotypes (e.g. Masters, 1953 Globigerina hoterivica Subbotina, p. 50,
1977, plates 25,26, and Michael, 1973, plate 5) that pI. 1, figs 1-4.
strict topotypes of F. washitensis can have four to 1964 Globigerina quadricamerata Antonova,
four and a half chambers visible dorsally in the last pp. 60, 61, pI. 12, figs 6a-c.
whorl, and three and a half to four chambers visible 1964 Globigerina tardita Antonova, p. 60, pI.
on the umbilical side. Longoria's proposal (1974) 12, figs 4-5.
for a second neotype was quite unnecessary, even 1986 Globuligerina hauterivica (Subbotina)
though it was useful in that a topotypic bullate (sic), Gorbachik, pI. 11, figs 1,2.
specimen was imaged. 1986 Globuligerina quadricamerata (Antonova),
The status of Reticuloglobigerina Reiss (1963) Gorbachik, pI. 12, figs 1-3.
was declared nomen nudum et oblitum by Loeblich 1986 Globuligerina tardita (Antonova), Gorba-
and Tappan, 1984, p. 38. chik, pI. 13, figs 1-3.
1988 Globuligerina hoterivica (Subbotina),
Range: Late Aptian-Early Cenomanian. The Ascoli, pI. 1, figs 14-16.
range of F. washitensis is well known from Early 1995 Favusella hoterivica (Subbotina), Wernli
Albian to 'Middle' Cenomanian, according to et al., pp. 379-390, pI. 1, figs 1-7; pI. 2,
Caron (1985), and it may also occur in horizons figs 1-16; pI. 3, figs 1-15.
as old as Late Aptian (Grigelis and Gorbachik,
1980). F. confusa was described from the Late Remarks: F. hoterivica has a small test; four
Albian of the Sierra de Papagayos, northeast of chambers in the final whorl; an intra-umbilical
Monterrey, Nuevo Leon State, northeastern aperture; the surface is finely reticulate (unlike the
Mexico (Longoria and Gamper, 1977). broadly reticulate F. washitensis (Carsey.
 Systematics 61

Range: Early Berriasian-Early Aptian. This and North West Africa, dated as Late Hauterivian
species was first described from the Hauterivian on the occurrence of nannofossils. Similar speci-
of northern Caucasus, and it was also recorded as mens were figured as Hedbergella hoterivica by
an index species from the Early Hauterivian of Butt (1979) from the Late Hauterivian-Barremian
Azerbaijan. Similar specimens were recorded as of the same site in the eastern North Atlantic.
'G.' tardita from the Late Barremian-Early Aptian
Ascoliella Banner and Desai, 1988
of the northwestern Caucasus by Antonova (in
Antonova et at., 1964), and by Gorbachik (1986), Type species Ascoliella scotiensis Banner and
and as 'G.' quadricamerata from the Early Desai, 1988 = Favusella nitida Michael, 1973
Aptian of northwestern Caucasus by Antonova (in
Banner and Desai (1988) erected a new genus
Antonova et al., 1964) and Gorbachik (1986).
Ascoliella for those forms with a favose reticula-
F. hoterivica, the oldest Favusella, was thought to
tion, like Favusella, but unlike the latter, having
have evolved in the Hauterivian from Cono-
an intra-extraumbilical aperture. Banner and Desai
globigerina (Banner and Desai, 1988, p. 148) and
(1988) designated a new species A. scotiensis as
was recorded as an index species for the Early
a type species for their new genus. However, it
Hauterivian of Azerbaijan by Ali-Zade and
was subsequently found that this species is iden-
Khalilov (1986). However, Wernli et al. (1995)
tical to F. nitida Michael, 1973 and should be its
found this form in the Berriasian and Valanginian
junior synonym.
of offshore Canada. Therefore, the continuous
reticulation occurred for the first time in the Ascoliella nitida (Michael, 1973)
Berriasian and the range of F. hoterivica must be
(Plate 4.5, Figs 1-9; Plate 4.7, Figs 1-3; Figure
emended. The breadth of the cells of the reticulum
4.1)
appears to remain significantly unchanged from
the Berriasian to Early Aptian. 1973 Favusella nitida Michael, p. 214, pI. 3, figs
10-12.
Favusella sti/tia RosIer, Lutze and Ptlaumann,
1973 Favusella pessagnoi Michael, pp. 214,
1979 215, pI. 4, figs 4--6.
(Plate 4.3, Figs 1-3; Figure 4.1) 1976 Favusella aff. washitensis (Carsey),
Ascoli, p. 661,674, pI. 1, figs 4a--c.
1979 Favusella stiftia RosIer, Lutze and 1977 Favusella hedbergellaeformis Longoria
Ptlaumann, p. 276, pl.1, figs 2-4. and Gamper, p. 207, pI. 4, figs 1-3,7-9.
1979 Hedbergella hoterivica (Subbotina), Butt, 1988 Ascoliella scotiensis Banner and Desai, p.
pI. 3, figs 1-4. 150, pI. 2, figs 3-4.

Remarks: This species has four chambers in the Remarks: This species has five to six subglob-
final whorl, the last one being reduced in size; ular chambers in the last whorl; a small and deep
surface reticulations are average in size, larger umbilicus; an intra-extraumbilical aperture; and a
than those of F. hoterivica (Subbotina) but smaller surface covered by anastomosing ridges which
than those of F. washitensis (Carsey); the reticu- enclose groups of microperforations in a medium
lation ridges are much more regular than in size of favose reticulation.
F. hoterivica. The aperture is intraumbilical, In 1973 Michael described F. pessagnoi from
covered by a bulla; the bulla carries a weak retic- the Late Albian of the very same locality as A.
ulation, with cells smaller than those of the retic- nitida. In the former, each cell of the reticulum is
ulation covering the chamber wall. smaller than in other forms of A. nitida, but these
species are otherwise identical. Banner and Desai
Range: Hauterivian-Barremian. This species (1988) designated as their holotype for Ascoliella
was first described from the DSDP Site 397, scotiensis the specimen figured by Ascoli (1976)
eastern North Atlantic, between the Canary Islands from the Aptian of the Oneida Well 0-25, side
62 The FavuseUidae

wall core at 7605 feet; this specimen is identical and obscuring part of the umbilicus; the umbilicus
to A. nitida. Longoria and Gamper (1977) figured is wide and deep; the aperture is intra-extra-
similar specimens to A. nitida, calling them F. umbilical; the surface of the test is covered with
hedbergellaeformis, from the Late Albian of the a medium-size favose reticulation.
Sierra de Papagayos, northeast of Monterey,
Nuevo Leon State, northeastern Mexico; we regard Range: Albian. F. scitula was first described
them as conspecific. from the Late Albian, Duck Creek Formation
(Washita Group) and from the overlying Fort
Range: Late Aptian-Late Albian. This species Worth and Denton formations of Texas. Longoria
was first described from the Late Albian of the and Gamper (1977) recorded this species as F.
Duck Creek, Fort Worth and Denton formations papagayosensis from the Albian of the Sabinas
of north central Texas, and subsequent records of Basin of northern Mexico.
synonymous forms show that the species has a
Late Aptian to Late Albian range. A. nitida is Ascoliella voloshinae (Longoria and Gamper,
the stratigraphically oldest known species of 1977)
Ascoliella. (plate 4.7, Figs 7-9; Figure 4.1)
Ascoliella quadrata (Michael, 1973) 1977 Favusella voloshinae Longoria and
(Plate 4.5, Figs 10-12; Figure 4.1) Gamper, p. 208, pI. 3, figs 1-3, 16-18.

1973 Favusella quadrata Michael, p. 215, pI. 4, Remarks: This species has four to five cham-
figs 7-9. bers ventrally, and the last chamber is often
smaller than the penultimate one; the umbilicus is
Remarks: This species has five chambers in the narrow and deep with an intra-extraumbilical aper-
last whorl ventrally, increasing gradually in size; ture; the surface of the test is covered with a very
the umbilicus is wide and shallow; the aperture is coarse favose reticulation.
intra-extraumbilical; the surface has a fine favose
reticulation.
Range: Late Albian. A. voloshinae was first
F. quadrata differs from F. nitida (Michael) in
described from the Sierra de Papagayos, northeast
having a much wider umbilicus, more globular
of Monterrey, Nuevo Leon State, northeastern
chambers and a much finer favose pattern.
Mexico.
Range: Late Albian. A. quadrata was first
described from the Duck Creek Formation
4.3 CONCLUDING COMMENTS
(Washita Group), and it was also found in the
overlying Fort Worth and Denton formations of
It was pointed out by Gorbachik and Kuznetsova
Texas.
(1986) that Globuligerina, Conoglobigerina and
Ascoliella scitula (Michael, 1973) Favusella all possessed walls which were pri-
marily aragonitic (Chapter 1). This mineralogy
(Plate 4.6, Figs 1-3; 8-10; Figure 4.1)
would be unique within the Globigerinina and
1973 Favusella scitula Michael, p. 215, pI. 4, alone would require separation of the genera into
figs 10-12. a distinctive superfamily. This distinction is
1977 Favusella papagayosensis Longoria and supported by the unique pseudomuricae and the
Gamper, p. 207, pI. 4, figs 16-18 (holo- favose reticulation in the Favusellidae, which has
type), 19-21; pI. 5, figs 16-21 (paratypes). no parallel elsewhere in the Globigerinina.
The earliest known true Favusella is F. hoteri-
Remarks: This species has five globular cham- vica ('Globigerina' hoterivica Subbotina, 1953,
bers in the last whorl, the last chamber protruding Hauterivian, Caucasus). Although this species was
 Concluding comments 63

recorded as an index fossil for the Early to 20 or more microperforations in each space
Hauterivian of Azerbaijan by Ali-Zade and (Plate 4.4, Fig. 8). A. voloshinae (Longoria and
Khalilov (1986), it is now believed that it evolved Gamper) (Plate 4.7, Figs 7-9) from the Late Albian
from Conoglobigerina much earlier, in the of NE Mexico has a coarse reticulum. Most species
Berriasian and Valanginian of offshore Canada of Favusella (like the intraumbilical Cono-
according to Wernli et at. (1995). Topotypes globigerina) possess a bulla covering the umbili-
photographed by SEM (Plate 4.1 , Figs 1-5, herein, cus (Plate 4.3, Fig. 2, Plate 4.4, Figs 3, 4), often
and Gorbachik, 1986, pI. 11, figs 1-2) possess a only a trace of it would be seen (Plate 4.2, Fig. 2).
continuous reticulum but with narrow inner ridge Ascoliella appears to have evolved from
spaces (containing only two or three microper- Favusella in the Late Aptian by migration of the
forations). In Favusella stiftia (RosIer et at., 1979) anterior end of the aperture beyond the umbilicus,
from the Late Hauterivian-Barremian of DSDP and is therefore the only known example in the
Site 397 (Plate 4.3, Figs 1-2) the reticulum spaces Favusellacea of a parallel to the evolution of
are broader and contain clusters of about five to Praehedbergella from Gorbachikella in the
ten microperforations. From the Aptian, the inner- Praehedbergellidae, Globigerinacea. Ascoliella
ridge spaces of the reticulum have broadened and acquired the broadest inner space-ridges between
the majority are flat-floored, containing at least ten the reticulum in the Late Albian but the genus
microperforations. F. washitensis (Carsey) (Plate appears to become extinct before the Cenomanian,
4.4, Figs 1-9) has very broad inner-ridge spaces whereas Favusella persisted into the Early
of the reticulum, and these are flat-floored, with 15 Cenomanian.
Plate 4.1

Figures 1-17 Favusella hoterivica (Subbotina, 1953). 1-5, Hauterivian, northern Caucasus, Crimea, deposited in the All-Union
Petroleum Scientific Research Prospecting Institute (VNIGRI), St Petersburg, Russia, paratype: I, dorsal view
(x260); holotype: 2-5, enlargement of surface wall showing favose reticulations (xl 100), peripheral, dorsal and
ventral views (x275). 6, Valanginian, Oneida 0-25 Well, GSC no. 53768, Scotian Shelf, Atlantic Canada (from
Ascoli, 1976, pI. I, fig. 3b), ventral view (x200). 7-12, (= Globigerina tardita Antonova), Late Barremian-Early
Aptian, along the Tusheps River, northwestern Caucasus: 7-9, holotype (from Antonova, 1964, pI. 12, figs 4a-c),
deposited in the Krasnodar Branch of the All-Union Oil and Gas Geological-Prospecting Institute, Krasnodar,
Russia: dorsal, ventral and peripheral views (x170); 10-12, topotypes (from Gorbachik, 1986, pI. 13, figs la, b,
2): 10, ventral view (x300), 11, dorsal view (x220) and enlargement of the surface showing the favose reticula-
tion (x1500). 13-17, (= Globigerina quadricamerata Antonova), Early Aptian, along the Kura River, north-western
Caucasus: 13-15, holotype (from Antonova, 1964, pI. 12, figs 6a-c), deposited in Krasnodar Branch of the All-
Union Oil and Gas Geological-Prospecting Institute, Krasnodar, Russia: dorsal, ventral and peripheral views
(xI50); 16, 17, topotypes (from Gorbachik, 1986, pI. 12, figs la, 2a), ventral views (x225).
Plate 4.2

Figures 1-8 Favusella hoterivica (Subbotina, 1953). Berriasian-Early Valanginian, from offshore wells, Eastern Canada (from
Wernli, Ascoli and Williams, 1995, pI. 2, figs 3, 5; pI. 3, figs 5, 13, 14; pI. 4, figs 5, 6, 8). Early Valanginian:
1, 2, ventral views (x260). Berriasian: 3, ventral view of specimen with bulla (x260); 4, dorsal view (x260);
5, peripheral view (x260); 6, 7, details of the irregularly distributed microperforations between the reticulum
(x 1000); 8, broken section of the (recrystallized) canaliculate wall (x1300).
Plate 4.3

Figures 1-3 Favusella stiftia RosIer, Lutze and Pfiaumann, 1979. Late Hauterivian, sample 397A-47-1, 5-8 em, DSDP Site
397, eastern North Atlantic, between the Canary Islands and NW Africa, holotype deposited in the collections of
the Geological-Palaeontological Institute and Museum, University of Kiel, Germany (from RosIer et al., 1979,
pI. I, figs la-<:): dorsal, umbilical (showing a bulla covering the umbilicus), and peripheral views (x41O).
Figures 4-6 Favusella orbiculata Michael, 1973 (nomen dubium) , a deformed specimen of Favusella, Late Albian, Weno
Formation, Washita Group, Texas, holotype deposited in the US National Museum of Natural History, Washington,
DC, no. 172795 (from Michael, 1972, pI. 4, figs 1-3): ventral, and dorsal views (x21O).
Figures 7-10 Favusella washitensis (Carsey, 1926) (=Favusella conJusa Longoria and Gamper, 1977), Late Albian, northern
Mexico, deposited in the US National Museum of Natural History, Washington, DC (from Longoria and Gamper,
1977, pI. 3, fig. 12; pI. 4, figs 25-27): 7-9, holotype (no. 207238), dorsal, peripheral and ventral views; 10,
paratype, ventral view (magnifications not given).
Plate 4.4

Figures 1-9 Favusella washitensis (Carsey, 1926). 1-4, Early Cenomanian, Street Bridge, Austin, Texas, USA, neotype (from
Longoria, 1974, pI. 26, figs 4-7), deposited in the US National Museum of Natural History, Washington, DC, no.
185021: dorsal, ventral and peripheral views (x75), and enlargement of the bulla covering the umbilicus (xI70).
5-9, Early Cenomanian, Paw Paw Formation, Washita Group, Grayson County, Texas, USA, deposited in The
Natural History Museum, London, nos P 52138, P 52139: 5, peripheral view (xI05); 6, ventral view (x115);
7, enlargement of same specimen (xI95); 8, enlargement of surface to show reticulations with microperforations
(x650); 9, enlargement of the umbilicus (x250).
Plate 45

Figures 1-9 Ascoliella nitida (Michael, 1973). 1-3, (= Ascoliella scotiensis Banner and Desai, 1988); form with small and
deep umbilicus (from Ascoli, 1976, pI. 1, fig 4a-c), Aptian, Oneida Well 0-25, offshore Canada, specimen
deposited in the Geological Survey of Canada, no. 53770: dorsal, peripheral and ventral views (xI20). 4--6, Late
Albian, Duck Creek Formation, Washita Group, Grayson County, Texas (from Michael, 1973, pI. 3, figs 10-12),
holotype of Favusella nitida Michael, deposited in the US National Museum of Natural History, Washington,
DC, no. 172793: dorsal, peripheral and ventral views (x90). 7-9, (= Favusella pessagnoi Michael, 1973), Late
Albian, Washita Group, Fort Worth Formation, Denton County, Texas (from Michael, 1973, pI. 4, figs 4--6),
holotype deposited in the US National Museum of Natural History, Washington DC, no. 172797: dorsal, ventral
views and twisted ventral view to show the intra-extraumbilical aperture (xllO).
Figures 10-12 Ascoliella quadrata (Michael, 1973). Late Albian, Washita Group, Duck Creek Formation, Texas (from Michael,
1973, pI. 4, figs 7-9), holotype deposited in the US National Museum of Natural History, Washington, DC, no.
172799: ventral, peripheral and dorsal views (xl00).
Plate 4.6

Figures 1-3.8-10 Ascoliella scitula (Michael, 1973). 1-3, Late Albian, Washita Group, Duck Creek Formation, Fort Worth,
Texas (from Michael, 1973, pI. 4, figs 10-12), ho10type deposited in the United States National Museum,
Washington, DC, no. 172801: dorsal, peripheral and ventral views (x11O). 8-10, (= Favusella papagayosensis
Longoria and Gamper, 1977), Albian, Tamau1ipas Formation, Sierra de Papagayos, Mexico (from Longoria
and Gamper, 1977, pI. 5, figs 16--18), paratype deposited in the US National Museum of Natural History,
Washington, DC: dorsal, peripheral and ventral views (magnification not given).
Figures 4-7 A deformed ?Ascoliella (=Favusella wenoensis Michael, 1973, nomen dubium). (From Michael, 1973, pI.
5, figs 4, 6, 8; pI. 7, fig. 3.) 4, 5, 7, Various views (xI40); 6, detail of reticulum (x950).
Plate 4.7

Figures 1-3 Ascoliella nitida (Michael, 1973) (= Favusella hedbergellaeformis Longoria and Gamper, 1977). Late Albian, Lower
Cuesta del Cura Formation, Sierra de Papagayos, Mexico (from Longoria and Gamper, 1977, pI. 4, figs 1-3), holo-
type deposited in the US National Museum of Natural History, Washington, DC, no. 207239: dorsal, peripheral
and ventral views (magnification not given).
Figures 4-6 Favusella hiltermanni (Loeblich and Tappan, 1961). Early Cenomanian, Zeltberg Brickworks, Hamburg, Germany
(from Loeblich and Tappan, 1961, pI. 4, figs 13a-{:), holotype deposited in the US National Museum of Natural
History, Washington, DC: dorsal, peripheral and ventral views (><90).
Figures 7-9 Ascoliella voloshinae (Longoria and Gamper, 1977). Late Albian, northeastern Mexico (from Longoria and Gamper,
1977, pI. 3, figs 1-3), holotype deposited in the US National Museum of Natural History, Washington, DC, no.
207237: dorsal, peripheral and ventral views (magnification not given).
 5
The practical taxonomy
of the Praehedbergellidae
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

5.1 INTRODUCTION known, the characters of the test as a whole may

be used with much lower magnification - with an
The morphology of the praehedbergellid test is optical microscope alone.
very simple but its variation reflects the biological When large assemblages of the Praehed-
differences which distinguished species. For prac- bergellidae are studied, it may be found that some
tical purposes, the morphospecies may themselves morphogroups always remain distinct from others;
be gathered into groups which we may call there are no intergradations between them. Other
generic- and family-group taxa. morpho groups display continuous gradations in
It is important that the differences we use can differences of shape; the extremes of difference
be recognized with an optical microscope; stereo- cannot be separated but the morpho group as a
scan images may help us understand the reasons whole is distinct from any other morphogroup.
for these optical distinctions, but it is not practical Examples which will be illustrated in Chapters 7
to have to depend on them. In an analogy, we may and 8 are, respectively, Praehedbergella tatianae
investigate the nature of the building materials of Banner and Desai (p. 109), which retains its par-
our rooms by sampling them and subjecting them ticular shape from individual to individual,
to microscopic study; this may tell us that some and Blefuscuiana speetonensis Banner and Desai
materials are wood, some are metal and some (p. 132), which varies in the convexity, flatness or
are plastic. However, once we know the detailed concavity of its spiral side, this variation being
structure of these materials, we can recognize continuous between individuals. If we have never
them without a microscope - we can see the super- found intergradation between one morphogroup
ficial differences between wood, metal and plastic and another, then we rate them as being separate
without the preliminary microscopic study. It is species. If we have recognized the continuity of
similar in our studies of the Praehedbergellidae. variation within the morphogroup, we rate these
An SEM image may be needed to define and variations as being of sub specific rank if the vari-
measure the details of the microperforations, ation is relatively restricted in time or space or
perforation cones, etc., but once the details are both; if there is no such restriction, we do not
80 The practical taxonomy of the PraehedbergeUidae

consider these variations to be worthy of taxo- (c) the degree of intercameral sutural depression
nomic distinction. An example of these may well at the periphery (measured here by the angle
be the presence or absence of perforation cones; between the chambers, the 'intercameral angle
Blefuscuiana infracretacea (Glaessner) s.s. has of sutural depression', at the periphery);
well developed perforation cones and occurs in (d) the size and relative depth of the umbilicus
both east and west Europe, while B. infracretacea (the latter being determined primarily by the
occidentalis BouDagher-Fadel et ai. lacks per- degree of overhang of the chambers of the 1st
foration cones and is dominant in the assemblages whorl; it may be shallow or deep);
only of western Europe. The greatest range of (e) the number of chambers seen ventrally to
individual variation within a monospecific assem- actually abut the umbilicus in the last mature
blage known to us is in Wondersella athersuchi whorl;
Banner and Strank; the adult chambers vary (f) the shape of the aperture (e.g. slit-like or high-
greatly in the assemblage, which we ascribe to arched);
dysoxicity (Chapter 13). This is exceptional, as (g) the nature of the test-wall surface (whether or
almost all other taxa follow clear morphological not there are perforation cones).
limits.
We have endeavoured to comply with the The quality of these morpho characters can, with
International Code of Zoological Taxonomy (3rd experience, be recognized rapidly by optical
edn, 1985) in all cases; other terms (such as those stereomicroscopy in well preserved specimens.
used by Saito, Thompson and Breger, 1981, p. 22) Groups which differ only in a single character (e.g.
have some value although they are not all recog- the magnitude of the intercameral angle of sutural
nized by the ICZN. 'We must all speak the same depression) are usually considered to be of varietal
taxonomic language and obey these rules of that or subspecific status.
language if we are to understand each other and Only 'adult' tests can be used in this way; this
if our work is not to be in danger of being totally means that about three whorls of chambers should
confused and worthless' (Derek Ager, 4/4/1985, be present. The tests being compared should be of
in The New Scientist, p. 45). approximately similar size.
We have preferred to 'split' rather than 'lump', In order to introduce a scheme which, we
while recognizing that future studies may show believe, should make such morphogroup compari-
at least some of our species or subspecies to be sons precisely repeatable, we have endeavoured to
synonyms. It will be easier, in the future, to dispense with reliance on vaguely used adjectives,
synonymise rather than to separate; it would be a usage which has been too common in the past.
specially difficult to disentangle the 'lumped' In order to identify the size of the specimens being
records, biostratigraphic or regional or palaeo- described, we give a measurement (in micromil-
environmental, of forms which it is later recog- limetres, often called microns) of the greatest
nized should have been referred to separate taxa. breadth (sometimes called diameter),
from the periphery to the opposite periphery, of
a typical specimen. Such a specimen may be a
5.2 CRITERIA WITIDN THE primary type (e.g. holotype, paratype) or a secon-
SPECIES-GROUPS dary type (e.g. topotype, metatype). From then
We have used the following characters to distinguish on, the shapes of the test components are given
between species or subspecies of similar test size: numerically as proportions, by the ratios of the
dimensions of one character to another. This not
(a) the height of coiling, number and shape of the only avoids the vague use of words alone, but also
chambers in each of the whorls, as seen allows direct, numerical comparisons to be made
dorsally; (regardless of the absolute size of the specimens).
(b) the rate of increase in chamber size with This is shown in Figure 5.1 , which can be
growth (either rapid or gradual); explained as follows.
 Criteria within the species-groups 81

2
6

7
4
Figure 5.1 Measurements used as an aid to the identification of the Praehedbergellidae: 1-4, Dorsal view: I, s =sutural depression,
b/a = rate of the spire opening; 2, h = height of each chamber, L = length of each chamber, hlL = chamber's dorsal shape (e.g.
hlL = 1 subglobular, hlL > 1 chamber radially elongate, hlL < 1 chamber depressed); 3,4 height of the chamber: 3, L' > L" the
height is greatest posteriorly; 4, L' < L" the height is greatest anteriorly. 5, Ventral view: s = sutural depression, uh = umbilical
height. 6, 7, Peripheral view: 6, convex test; 7, planispiral test (t = thickness of the chambers; h = height of chambers; d = height
of the spire).

We use dorsal to label the more evolute side of equiangular) is given by the ratio of measurements
a spiral test and ventral for the more involute side. made radially from the proloculus to the periphery
We do not use the term spiral rather than dorsal of the first chamber of the last whorl (a) and to
because both sides may show spirality. Compar- the periphery of the last chamber of the test (b);
ably, the term umbilical rather than ventral loses the value of b/a will then be a record of the rate
its meaning when there are umbilici (true or false of spire opening, regardless of the differences of
umbilici) on both sides of the test. precise measurements between specimens of
In dorsal view (Figure 5.1, 1), the rate of different absolute size.
opening of the spire (which mayor may not be The height of each chamber (h), in dorsal view,
82 The practical taxonomy of the Praehedbergellidae

is the radial dimension from the spiral suture to the These species-groups are gathered together in
periphery (Figure 5.1,2). The length of each cham- ways which portray stages in the evolution of the
ber (L) is the distance from one intercameral suture family, and which enable ready recognition of
to the next, a measurement perpendicular to the the species themselves.
height (Figure 5.1, 2-4). If the dorsal view of a We believe that all the evidence indicates
chamber shows its maximum height to be in the that the stratigraphically earliest microperforate
middle of that chamber then the value of hlL gives Praehedbergellidae (Late Valanginian) were
the chamber's dorsal shape. If hlL == 1, the chamber trochospiral and gave rise to some trochospiral
is likely to be subglobular (Figure 5.1, 2); if hlL > macroperforate descendants (the Hedbergellidae).
1, the chamber is radially elongate, and if hlL < 1, These trochospiral praehedbergellids were also
then it is depressed. The value of these ratios give immediately ancestral to planispiral forms, which
the degree of elongation or depression of the cham- did not appear until the Barremian; these also
bers. Some chambers are higher posteriorly or ante- gave rise to planispiral macroperforate ascendants
riorly than they are medially; then the height of the (the Planomalinidae) (Figure 5.2). Because these
chamber can be drawn to divide the length trochospiral and planispiral groups of the Prae-
unequally (Figure 5.1, 3-4). If the height is great- hedbergellidae are readily recognizable and show
est in the posterior part of the chamber, then successive stages in the evolution with separate
L' > L" (Figure 5.1, 3). Conversely if the height is descendants, we consider them to be successive
greatest anteriorly, then L' < L" (Figure 5.1, 4). families, the Praehedbergellidae and the Schacko-
Therefore the shape of the chambers in dorsal view inidae respectively.
can be numerically defined. The trochospiral Praehedbergellidae contains
In ventral (umbilical) view (Figure 5.1, 5) the many species groups which are morphologically
radial height of each chamber must be measured distinct and which represent successive evolution-
from the umbilical margin of the chamber to its ary stages. The oldest form possesses an aperture
periphery (uh, umbilical height). The umbilicus which is intraumbilical but this is followed strati-
has a maximum breadth u, so that the ratio uluh graphically by groups of species in which the aper-
characterizes the taxon. ture becomes intra-extraumbilical. This difference
The degree of sutural depression may be characterizes groups of species which we can label
measured in either the ventral or the dorsal view with different generic names. The former (with an
as the angle s (Figure 5.1, 1, 5). intraumbilical aperture) we call Gorbachikella; the
In peripheral view, when the dorsal-ventral latter group (with an intra-extraumbilical aperture)
extent of the spiral axis may be seen, the thick- comprises many morphologically different forms
ness (t) of the chambers may be measured (Figure which themselves constitute successive evolution-
5.1, 6, 7). The ratio hit is easily determined in ary stages and which themselves can be labelled
such views. If the initial whorls of the trochospire as different genera. These include the species-
are strongly convex, they make the dorsal side of groups called Praehedbergella (with few chambers
the test very high (Figure 5.1, 6); this height (d) in each adult whorl, like the ancestor Gorbachi-
can readily provide the ratio dlt, which gives a kella), Blefuscuiana (with more chambers per
numerical measure of dorsal test convexity. whorl, the chambers being subglobular to
depressed), Lilliputianella (with radially elongated
chambers), and Wondersella (with adult strepto-
5.3 SUPRASPECIFIC CRITERIA IN THE spirality) .
PRAEHEDBERGELLIDAE The trochospiral tests gave rise to planispiral
ones (the Shackoinidae). Some possessed cham-
The genus-group and family-group criteria of the bers similar to those of the ancestor Praehed-
Praehedbergellidae must be assessed by recog- bergella, and we call them Blowiella; others
nizing the morphological characters common to radially elongated their chambers (Claviblowiella,
different groups of species. Leupoldina, Schackoina).
 ...... ...... ...... ......
...... CD
W N N N ...... :s::
N ...... ...... i\) Q)

VALANGINIAN HAUTERIVIAN BARREMIAN APTIAN ALBIAN

EARLY LATE EARLY LATE EARLY LATE EARLY LATE

Gorbachikella
'\'\. IQ)
..."tJ
Praehedbergella CD CD
\ \ " LiIIlputianelloideS
=::r
-'CD
\ \ a. a.
\ \ Q)U
Lilliputianella
\ \ CD CD
\ \ I ...
\ BlefuSCLana I CC
\ \
"r
\
\ .:::J:
\ \ (I) CD
Hedbergella =Q.
\ "r Q.C"
\
m(l)
(1)-'
\ cc
\ Planomalina ."'0
\ ,.... =ii'
\ I 2.:::::1
/ Alanlordella Q.O
\ ,..... m3
\ I (l)m
\
\ / Globigerinelloides en
\ ()
/
\ Blowiella J ::r
Q)
\ ()
\
'\. Claviblowiella
~
\ 0
_.
\ Leupoldina
::::J
_.
\ Schackoina a.
Q)
"r CD
Figure 5.2 Phylogeny of the Early Cretaceous Globigerinacea (Praehedbergellidae, Hedbergellidae, Planomalinidae and Schackoinidae). Chronostratigraphy and Absolute Time
Scale after Gradstein et al. (1995).
84 The practical taxonomy of the PraehedbergeIlidae

All these genera are fully diagnosed in the mental information (Chapter 13), their geographic
following chapters. They may be recognized occurrences can give clues as to the palaeo-
unambiguously, their usage restricts the number of ceanography of the time, and their successive
species with which new specimens are to be morphological differences reveal the phylogenetic
compared, their stratigraphic ranges provide a changes which occurred during the evolution of
broad biostratigraphy, their relative dominances in the family.
microfossil assemblages can give palaeoenviron-
 6
The earliest
Praehedbergellidae
- Gorbachikella
M. K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

6.1 INTRODUCTION the Caribbean), from the Late Valanginian to the

Early Aptian (Figure 6.1). Gorbachikella grandi-
In the Late Valanginian of North Africa several apertura has clearly five chambers in the early
planktonic foraminiferal species appeared which whorls and could be the immediate descendant
were morphologically different from all the of Conoglobigerina in the Late Valanginian. G.
Favusellacea of earlier times. The new taxa were anteroapertura also has an anteriorly facing aper-
equally microperforate but they were superficially ture and occurs in beds as old as Late Hauterivian;
smooth, lacking all muricae or surface ridges. All we believe that this modification of the aperture led
had four chambers in the last whorl and these to one with an intra-extraumbilical extent and
chambers were subglobular, being approximately evolved into the genus Praehedbergella and, sub-
as high as long and almost equally thick. The aper- sequently, all the remaining Praehedbergellidae.
ture was an intraumbilical arch, furnished with a Therefore, we believe that Gorbachikella was the
narrow rim-like lip, and the umbilicus itself was direct and immediate ancestor of all the Mesozoic
open and deep. Such forms have been named Globigerinacea.
Gorbachikella by Banner and Desai, 1988; similar Although most forms of Gorbachikella are
species were called Caucasella by Longoria, 1974, consistent in morphology and can be clearly
and many later authors, but this is a homonym recognized as distinct species, some are clearly
(Caucasella Moisseev, 1934) and its type species distorted (Plate 6.2, Figs 7-9); it is possible that
was Globigerina hoterivica Subbotina, 1953, the five specimens described by Maamouri and
which we now know to be a Favusella. Salaj, 1995, as 'Archaeokassabella biapertura
Different species of Gorbachikella may be found n. gen. and sp.' (Plate 6.2, Figs 10-11) are also
in the Early Cretaceous sediments of North Africa, distorted and are in fact a deformed species of
Eastern Europe and Central America (including Gorbachikella.
.A .A .A .A .A .A
Co) Co) I\,) I\,) I\,) a::
~ I\,) o .... CJ) .A I

VALANGINIAN HAUTERIVIAN BARREMIAN APTIAN

EARLY LATE EARLY LATE EARLY LATE EARLY

~
Gorbachikel/a anteroapertura
G.depressa
G. grandlapertura
G. kuglerl
G. neill

J J J L:. I
fTt P
Ii!
so
C'I
~r-
...
P
&.
~
- ~
& ICI"2-
f Nannofossil
& r fIi atI'== 'i
~ datums
Ii
~
'*
I II I ~
II I . I I I I
.,1 I~,",I I
I~I I ! b,1!1
. II .a:.
t
iii - c: lam - i~ Ammonite
~
i: I:' If l'~~' i 'I
-I r- ~
1:1 i- I' '*'~ Ii I Ii':" ~11.1 =. IS!: zones
I II I
- I II
.(' I I I I I

Figure 6.1 Stratigraphical distribution of Early Cretaceous Gorbachikella. Nannofossil datums and stratigraphic/biostratigraphic correlation based on Bergen (1994).
Chronostratigraphy and Absolute Time Scale after Gradstein et al. (1995).
 Systematics 87

6.2 KEY TO THE SPECIES OF Remarks: G. kugleri has four chambers in each
GORBACHIKELLA of its three whorls. The chambers are slightly
=
depressed (hiL 0.85 approx.), and subglobular
1 Aperture antero-intraumbilical (hit =1 approx.). The umbilicus is broad (u/uh =
G. anteroapertura 0.27 approx.) The semi-circular, axio-intraumbili-
2 Aperture axio-intraumbilical: cal aperture possesses a thin rim-like lip. The
2.1 Aperture with a thin rim, umbilicus wide: dorsal side is virtually flat.
2.1' Aperture broad and high
G. grandiapertura Range: Hauterivian-Early Aptian. Gorbachi-
2.1" Aperture a semicircular arch kella kugleri was originally described (Bolli, 1959,
G. kugleri p. 270) from the Barremian of Trinidad, West
2.1'" Aperture a low opening Indies. Marianos and Zingula, 1966, reported it
G. depressa from the Hauterivian-Barremian of California,
2.2 Aperture with a broad lip, umbilicus USA, while Longoria, 1974, illustrated it (as
narrow G. neili Caucasella hoterivica) from the earliest Late
Aptian of Mexico. BouDagher-Fadel et al., 1995,
found it to range from the Hauterivian to the Early
6.3 SYSTEMATICS Aptian in Tunisia. In addition, we have for the
first time identified this species from the Early
Superfamily GLOBIGERINACEA Carpenter, Barremian of Kurovice, Magura Flysch, West
Parker and Jones, 1862 Carpathians (sample 3H, collected and donated by
Dr M. Bubik). It has not been found in the
Family PRAEHEDBERGELLIDAE Banner Barremian-Aptian of the North Sea (Banner et al.,
and Desai, 1988 1993).
Gorbachikella anteroapertura BouDagher-Fadel
Subfamily PRAEHEDBERGELLINAE subfam.
et al., 1995
nov.
(Plate 6.3, Figs 1-10; Figure 6.1)
Genus Gorbachikella Banner and Desai, 1988
1981 Caucasella hoterivica (Subbotina), Bellier
Type species Globigerina kugleri Bolli, 1959
and Chitta, pI. 1, figs 1-2.
Gorbachikella kugleri (Bolli, 1959) 1995 Gorbachikella anteroapertura BouDagher-
Fadel et al., p. 190, pI. 5, figs 1-6.
(Plate 6.1, Figs 1-6, Plate 6.2, Figs 1-6; Figure
6.1)
Remarks: G. anteroapertura has four chambers
1959 Globigerina kugleri Bolli, p. 270, pI. 23, in each of its three whorls. The chambers are
figs 3-5. slightly depressed (hiL =0.78 to 0.80), being
?1966 Globigerina kugleri Bolli; Marianos and slightly more depressed than in G. kugleri. The
Zingula, p. 335, pI. 37, figs 4a-c. umbilicus is fairly broad (u/uh =0.19 approx.).
1974 Caucasella hoterivica (Subbotina), The semi-circular, anteroumbilical aperture
Longoria, p. 49, pI. 11, figs 9-11, 14-16. possesses a narrow rim-like lip. The dorsal side is
1988 Gorbachikella kugleri (Bolli), Banner and slightly convex.
Desai, p. 151, pI. 2, fig. 2.
?1991 Globuligerina hoterivica (Subbotina), Range: Late Hauterivian-Early Aptian. G.
Altiner, pI. 14, figs 1-7. anteroapertura was reported and illustrated (as
1995 Gorbachikella kugleri (Bolli), BouDagher- Caucasella hoterivica) from the Late Barremian
Fadel et al., p. 188, pI. 2, figs 1-5. and Early Aptian of Jebel Goraa in Tunisia by
Bellier and Chitta, 1981. BouDagher-Fadel et al.,
88 The earliest Praehedbergellidae - Gorbachikella

1995, recorded it as ranging from the Late Remarks: G. grandiapertura has a very high
Hauterivian to the Early Aptian of Tunisia. In and broad axio-intraumbilical aperture, five cham-
addition to its occurrence in North Africa, we have bers in the first whorl and four in each subsequent
also found this species in sample 3H (Early whorl, and a high spiral side (dlt = 0.29 approx.).
Barremian), collected by Dr M. Bubik from As we have not seen actual specimens of G.
Kurovice, West Carpathians (Magura Flysch). compressa, and as the type specimen was inade-
quately figured (there was no umbilical view), we
Gorbachikella depressa BouDagher-Fadel et al.,
cannot be sure of its taxonomy and have only
1995
doubtfully placed it here. G. spiralis, published in
(Plate 6.4, Figs 1-9, Plate 6.6, Figs 2-3; Figure the same paper by Maamouri and Salaj, is,
6.1) however, clearly synonymous.
1984 Caucasella hoterivica (Subbotina), Salaj,
pI. 1, fig. 7-9. Range: This species in Tunisia ranges from Late
1987 Caucasella hoterivica (Subbotina), Ben Valanginian to Late Barremian (BouDagher-Fadel
Haj Ali, pI. 1, figs 13-15. et al. 1995; Maamouri and Salaj, 1995).
1995 Gorbachikella depressa BouDagher-Fadel Gorbachikella neili (Maamouri and Salaj, 1995)
et al., p. 188-190, pI. 3, figs 1-4.
1995 Globuligerina mejezensis Maamouri and (Plate 6.6, Figs 7-8; Figure 6.1)
Salaj,pp.132,p1.3,figs.1,2,pI.5,figs 1-6. 1995 Globuligerina neili Maamouri and Salaj,
pp. 131, 132, pI. 1, figs 1-5.
Remarks: Another species previously mistaken
for Globigerina hoterivica Subbotina, 1953. G. Remarks: The test has a very low axio-intra-
depressa has an axioumbilical aperture and four umbilical aperture with a broad lip throughout its
chambers in each whorl; it differs from G. kugleri length; the umbilicus is narrow and the chambers
in having more depressed chambers (hIL= 0.67 are depressed (uluh = 0.29 approx.; hlL = 0.6
approx.), a broader intercameral sutural angle, a approx.); there are four chambers in each of the
very low, symmetrical aperture, and a high spire three whorls.
(dlt = 0.26 approx.).
Range: This species is known from the Late
Range: Gorbachikella depressa is presently
Valanginian and Early Barremian of Tunisia.
known only from Tunisia, where it ranges
throughout the Late Valanginian to the Late
Barremian.
6.4 CONCLUDING REMARKS
Gorbachikella grandiapertura BouDagher-Fadel
et al., 1995 After many years of neglect, two articles on the
first and most primitive praehedebergellid,
(Plate 6.5, Figs 1-9, Plate 6.6, Figs 1,4-6;
Gorbachikella, appeared in the same year
Figure 6.1)
(BouDagher-Fadel et al., in September, 1995, and
1995 Gorbachikella grandiapertura BouDagher- Maamouri and Salaj, in November, 1995). Both
Fadel et al., p. 190, pI. 4, figs 1-5 articles are on the Early Cretaceous of Tunisia;
[published September]. BouDagher-Fadel et al. dealt with the Hauterivian,
?1995 Globuligerina compressa Maamouri and Barremian-Aptian planktonic foraminifera,
Salaj, p. 133, pI. 6, figs 1-2 [published Maamouri and Salaj with those of the latest
November]. Valanginian. As BouDagher-Fadel et al.'s article
1995 Globuligerina spiralis Maamouri and appeared before that of Maamouri and Salaj,
Salaj, p. 132, pI. 2, figs 1-4, pI. 3, figs 3--6, where there are cases of synonymy, the former
pI. 4, figs 1-6 [published November]. must have priority.
 Concluding remarks 89

The planktonic assemblages of the Early surface of the wall is still microperforate, it lacks
Cretaceous of Tunisia were made up entirely of muricae.We believe that Gorbachikella is the
species of Gorbachikella. Only two species persist direct descendant of Conoglobigerina and is the
throughout and into the Aptian, where they are immediate ancestor of the first Praehedbergella.
joined by new forms of praehedbergellids. Gorbachikella has a very restricted geographic
BouDagher-Fadel et al. (1995) demonstrated that distribution (BouDagher-Fadel, 1995, p. 140,
Gorbachikella is a primitive species belonging to Figure 1).
the Praehedbergellidae and that although the
Plate 6.1

Figures 1-6 Gorbachikella kugleri (Bolli, 1959). The type-species with an intraumbilical, low symmetrical aperture. 1, 2, Early
Barremian, 5594, Beauvoir-I Well, Tunisia, deposited in The Natural History Museum, London, no. PF 52989:
ventral and dorsal views (x2554); 3-6, Late Hauterivian, 5574, Beauvoir-I Well, Tunisia, deposited in The Natural
History Museum, London, no. PF 52986: 3, 4, ventral and dorsal views of the same specimen as in figs 5 and 6,
but following an ultrasound washing in sodium hexametaphosphate (x245); 5, 6, ventral and dorsal sides before
cleaning (x235).
Plate 6.2

Figures 1-6 Gorbachikella kugleri (Bolli, 1959). 1-3, Barremian, Cuche Formation, 7102-7263 feet, Montesserat-l Well,
Trinidad (from Bolli, 1959, pI. 23, figs 3a-c), holotype deposited in the US National Museum of Natural History,
Washington, DC, no. 626289: dorsal, peripheral and ventral views (xllO). 4-6, Barremian, SS94, Beauvoir-I
Well, Tunisia, deposited in The Natural History Museum, London, no. PF 52988: 4, 6, dorsal and ventral views
(x230); 5, peripheral view showing clearly the axio-intraumbilical aperture of semi-circular shape with a thin
rim (x205).
Figures 7-9 Gorbachikella sp., deformed specimens in which the aperture has moved to the periphery and the umbilicus is
closed. Late Hauterivian, SS74, Beauvoir-I Well, Tunisia, deposited in The Natural History Museum, London,
nos PF 62947, PF 62948: 7, 8, dorsal and ventral views (x175); 9, dorsal view (x230).
Figures 10, 11 Archaeokassabella biapertura Maamouri and Salaj, 1995. Probably deformed specimens of Gorbachikella (from
Maamouri and Salaj, 1995, pI. 6, figs 5, 6). Late Valanginian, Jebel Oust, Tunisia; holotype deposited in collec-
tions of the National Geological Survey of Tunisia, no. 0-6: peripheral views (x205 and x225, respectively).
Plate 6.3

Figures 1-10 Gorbachikella anteroapertura BouDagher-Fadel et ai., 1995. A species with a slightly low, intraumbilical aper-
ture facing anteriorly, and an almost flat dorsal side. All specimens deposited in The Natural History Museum,
London, PF 52999 (holotype), PF 53000-PF 53004 (paratypes). Early Barremian, SS94, Beauvoir-I Well, Tunisia:
1, Holotype, ventral view showing the microperforations (x200); 2, paratype, ventral view (x270); 3, paratype,
ventral view showing an aperture nearly reaching the periphery (x175); 4, 5, paratype, ventral and dorsal views
(x250); 6, paratype, dorsal view (x220). Early Barremian, SS101, Beauvoir-I Well, Tunisia: 7, paratype, ventral
view (x250); 8-10, paratype, dorsal, peripheral and umbilical views (x220).
Plate 6.4

Figures 1-9 Gorbachikella depressa BouDagher et al., 1995. A species with a low symmetrical aperture and a convex spiral
side. All specimens deposited in The Natural History Museum, London, nos PF 52990 (holotype), PF 52991-
PF 52993 (paratypes). Early Barremian, SS638, Jebel Oust-III Well, Tunisia: I, 2, holotype, ventral and dorsal
views (x215). Early Barremian, SS101, Beauvoir-I Well, Tunisia: 3,5,6 paratype, ventral view showing the low
aperture, peripheral view showing a very convex dorsal side, and the dorsal view (xI85). Early Barremian, SS94,
Beauvoir-I Well, Tunisia: 4, paratype, ventral view (x220). Early Barremian, SS638, Jebel Oust-III Well, Tunisia:
7-9, paratype with a deformed last chamber, but showing clearly the low aperture and the microperforate test: 7,
8, dorsal and ventral views (x240); 9, enlargement of the surface of the last chamber showing the irregular micro-
perforations (xI250).
Plate 6.5

Figures 1-9 Gorbachikella grandiapertura BouDagher-Fadel et al., 1995. A species with an intraumbilical aperture (a high
symmetrical to slightly asymmetrical arch) and a highly convex dorsal side. All specimens deposited in The Natural
History Museum, London, nos. PF 52994 (holotype), PF 52995-PF 52998 (paratypes). 1-3, Early Barremian, SS94,
Beauvoir-I Well, Tunisia: holotype, ventral view (with a high aperture), dorsal view and peripheral view showing
a highly convex dorsal side (x205). 4, 5, Early Barremian, SS93, Beauvoir-I Well, Tunisia: paratype, ventral side
with a high aperture and very convex dorsal side (x230). 6, Early Barremian, SS94, Beauvoir-I Well, Tunisia:
paratype, ventral view (x250). 7, 8, Early Barremian, SSIOI, Beauvoir-I Well, Tunisia: paratype, ventral and dorsal
views (x255). 9, Early Barremian, SSIOI, Beauvoir-I Well, Tunisia: paratype, ventral view (x205).
Plate 6.6

Figures I ,4-6 Gorbachikella grandiapertura BouDagher-Fadel et al., 1995. I, (= Globuligerina compressa Maamouri and Salaj,
1995), Late Valanginian, Jebel Oust, holotype deposited in collections of the National Geological Survey of Tunisia,
Tunis (from Maamouri and Salaj, 1995, pI. 6, fig. 1): dorsal view (x260). 4-6, (= Globuligerina spiralis Maamouri
and Salaj, 1995), latest Valanginian, Jebel Oust; paratype and holotype deposited in the collections of the National
Geological Survey of Tunisia, Tunis, nos 0-2 and 0-3 (from Maamouri and Salaj, 1995, pI. 3, fig. 3; pI. 2, figs
1, 2): 4, ventral side (x250); 5, 6, dorsal and tilted ventral views (x340).

Figures 2-3 Gorbachikella depressa BouDagher-Fadel et al., 1995. 2, 3, (= Globuligerina mejezensis Maamouri and Salaj,
1995), Late Valanginian, Jebel Oust, holotype deposited in collections of the National Geological Survey of Tunisia,
Tunis, no. 0-4 (from Maamouri and Salaj, 1995, pI. 3, figs 1,2): dorsal and peripheral views (x31O).
Figures 7-8 Gorbachikella neili (Maamouri and Salaj, 1995). 7, Late Valanginian, Jebel Oust, deposited in the National
Geological Survey of Tunisia, Tunis, no. 0-1 (from Maamouri and Salaj, 1995, pI. 1, fig. 3): ventral side (x265).
8, Early Barremian, SS94, Beauvoir-I Well, Tunisia: ventral view showing the narrow umbilicus and the thick lip
(x250).
 7
The flowering of the
Praehedbergellidae:
Praehedbergella
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

7.1 INTRODUCTION Praehedbergella is microperforate and smooth and

of Early Cretaceous (Aptian and older) age only.
Unlike Gorbachikella, Praehedbergella had a Praehedbergella was redefined by Banner and
wide geographic distribution. It spread from low Desai (1988) and that redefinition was emended
to middle latitudes. It ranged in age from the Early again by Banner et al. (1993). They distinguished
Hauterivian to the Late Aptian (Figure 7.1) and Praehedbergella from Blefuscuiana by the pres-
its species have been recorded from localities as ence of only four chambers on the ventral side
far west as northern Mexico (Longoria, 1974), as abutting the umbilicus. The emendation was
far north as North Sea area (Banner et al. 1993) necessary as these forms are restricted only to the
and North Yorkshire, England (Banner and Desai Early Cretaceous, while those with more than four
(1988), and as far south as the Walvis Ridge, such chambers are known to range from the Early
South Atlantic (Caron, 1985). Barremian (e.g. Blefuscuiana praesimilis = B.
The original definition of Praehedbergella by laculata Banner et al. (1993), Plate 8.9, figs 1-3)
Gorbachik and Moullade (1973) was interesting to the Early Danian (e.g. B. archeocompressa
but very difficult to use. Those authors considered (Blow, 1979)). The holotype of Praehedbergella
Praehedbergella to be a subgenus of Clavihed- tuschepsensis (Antonova), the type species of
bergella Banner and Blow (1959) and to differ Praehedbergella was only drawn by Antonova
from it by its 'globigeriniform' (not digitiform) (1964), as figured here (Plate 7.1, Figs 1-3). It
chambers, and by its thinner wall (of 'mono- was deposited in the VNIGRI collection in St
lamellar structure', as compared to the 'bilamellar Petersburg, and for the first time, it has been here
structure' of Hedbergella). We now know that (Plate 7.1, Figs 4--7) photographed under the scan-
both Hedbergella and Clavihedbergella are ning electronic microscope (SEM). This has
macroperforate and muricate, and that both are of confirmed the validity of the emendation of this
Late Aptian and younger Cretaceous age, while genus by Banner et al. (1993). On the other hand,
....
Co) i:
.., ..,.......... ..,........
m

HAUTERIVIAN BARREMIAN APTIAN :t-

el
m
EARLY LATE EARLY LATE EARLY LATE
,

Praehedbergella handousl
P. pseudoslgall
P. ruka s.s.
P. ruka contrltus
P. ruka paplliata
P. slgall s.s.
P. slgall compacta
P. tatlanae
P. tuschepsensls s.s.
P. tuschepsensls grlgellsl
P. tuschepsensls perforare
P. yakovlevae

Figure 7.1 Stratigraphical distribution of Early Cretaceous Praehedbergella. Chronostratigraphy and Absolute Time Scale after Gradstein et al. (1995).
 Systematics 105

two paratypes deposited in the Moscow State 23.2 Four to five chambers in first whorl, final
University appear different from the holotype. As aperture low:
seen here (Plate 8.12, Figs 7-9), one paratype (that 23.2.1 Early chambers of last whorl not
illustrated dorsally by Gorbachik, 1986, plate 25, depressed and intercameral sutural angles rela-
fig. 1) is a young Blefuscuiana, while the other is tively small P. sigali s.s.
a different species of Praehedbergella (as yet 23.2.2 Early chambers of last whorl slightly
known only by this single specimen). Therefore, depressed and intercameral sutural angles rela-
the characters not only of P. tuschepsensis, but tively broad
also those of the genus Praehedbergella and of P. sigali compacta
the family groups based upon it, depend on the 2.4 Final chambers highest posteriorly
holotype in St Petersburg. We sincerely thank Dr P. yakovlevae sp.nov.
Svetlana Yakovleva for allowing us to borrow it
and re-illustrate it. 73 SYSTEMATICS

Superfamily GLOBIGERINACEA Carpenter,

Parker and Jones, 1862
7.2 KEY TO THE SPECIES OF
PRAEHEDBERGELLA Family PRAEHEDBERGELLIDAE Banner
and Desai, 1988
1 Dorsal side strongly convex, spire high
The Praehedbergellidae comprise those micro-
1.1 Slow rate of chamber enlargement
perforate, non-muricate genera which are trocho-
P. ruka s.s.
spiral and which have intraumbilical or umbilical-
1.2 More rapid rate of chamber enlargement:
extraumbilical ventral apertures.
1.2.1 Surface smooth P. ruka contritus
1.2.2 With perforation cones Genus Praehedbergella Gorbachik and Moullade,
P. ruka papillata 1973 (emend. Banner, Copestake and White,
2 Dorsal side weakly concave or weakly convex, 1993)
spire low
Type species Globigerina tuschepsensis
2.1 Chambers strongly depressed throughout,
Antonova, 1964
intercameral sutural angle very broad
P. tatianae Praehedbergella tuschepsensis (Antonova, 1964)
2.2 Chambers of last whorl slightly depressed sensu stricto emend.
throughout:
(Plate 7.1, Figs 1-7; Figure 7.1)
2.2.1 Test dorso-ventrally compressed
P. handousi 1964 Globigerina tuschepsensis Antonova,
2.2.2 Test dorso-ventrally compressed: pp. 59, 60, pI. 12, figs 3a-c.
2.2.2.1 Dorsal surface fiat, umbilicus broad: non 1986 Clavihedbergella tuschepsensis
2.2.2.1.1 Surface smooth (Antonova), Gorbachik, pp. 118, 119,
P. tuschepsensis s.s. 236, pI. 25, fig. 1.
2.2.2.1.2 With perforation cones non 1993 Praehedbergella tuschepsensis
P. tuschepsensis perforare (Antonova), Banner, Copestake and
2.2.2.2 Dorsal side slightly convex, umbilicus White, pp. 8, 9, pI. 2, figs 4a-c.
narrow
P. tuschepsensis grigelisi Emended diagnosis: Test microperforate and
23 Final chambers subglobular: smooth (no muricae or pustules of any sort: perfor-
23.1 Six and a half chambers in the first whorl, ation cones occur only in one subspecies). Four
final aperture high and a half chambers are visible dorsally in the last
P. pseudosigali whorl, but only four abut the ventral umbilicus.
106 The flowering of the Praehedbergellidae

The chambers of the last two whorls enlarge from the umbilicus to the periphery and is fur-
rapidly (b/a = 1.6 approx.) and the chambers of nished with a porticus. The spiral side is slightly
the last whorl are subglobular, being only very convex.
slightly depressed (h/L = 0.71 approx.) and highest The revision of the nature of this species shows
in their mid-points (L' = L"). The dorsal surface is us that P. tuschepsensis grigelisi differs from
virtually flat. The intra-extraumbilical aperture P. tuschepsensis s.s in possessing higher cham-
is a low slit, extending to the periphery of the pen- bers, a slightly more convex spiral side and a
ultimate whorl, and possessing a thin, narrow, narrower umbilicus.
lip-like porticus. The umbilicus is quite broad and
deep (u/uh = 0.24).
Range: Late Aptian. P. tuschepsensis grigelisi
was originally described from the Late Aptian
Remarks: P. tuschepsensis s.s. differs from its of Speeton Cliffs, Filey Bay, North Yorkshire,
subspecies P. tuschepsensis perforare Banner et Schackoina cabri Zone.
al. in lacking perforation cones and from the
subspecies P. tuschepsensis grigelisi Banner and Praehedbergella tuschepsensis (Antonova)
Desai in its flat dorsal surface and broader perforare Banner, Copestake and White, 1993
umbilicus.
(Plate 7.1, Figs 8-11; Figure 7.1)
As noted in the Introduction to this chapter, the
specimens given this species name by Gorbachik 1993 Praehedbergella grigelisi Banner and
(1986) and by Banner et al. (1993) were based Desai perforare; Banner, Copestake and
erroneously on the dorsal morphology of a White, p. 5, pI. 1, figs 1a-d.
paratype in the collection of the Moscow State
University; that specimen is now known to belong Remarks: P. tuschepsensis perforare has about
to a species (and genus) different from the holo- five chambers in the penultimate whorl, reducing
type of P. tuschepsensis s.s.
to four in the last whorl. The spiral side is almost
flat. The chambers are symmetrical and highest in
Range: The actual range of P. tuschepsensis s.s. their mid-points (L' =L") and increase signifi-
is not yet fully known; the holotype was obtained cantly in size and growth. The last chambers
from the Early Barremian, Spiroplectammina are slightly depressed (h/L = 0.92 approx.). The
magna beds, Tuscheps River (tributary of the umbilicus is shallow and broad. The aperture is a
River Pshish), northwest Caucasus. narrow slit bordered with a porticus. Perforation
Praehedbergella tuschepsensis (Antonova) cones are visible at least on the chambers of the
grigelisi Banner and Desai, 1988 last whorl.
Following the revision of P. tuschepsensis, this
(Plate 7.1 Figs 12-14; Figure 7.1) taxon (previously regarded as a subspecies of P.
1959 Globigerina sp. 1 Obregon de la Parra, grigelisi) is transferred to Antonova's species. P.
p. 150, pI. 3, figs 7a-b. tuschepsensis perforare differs from P. tuschep-
1988 Praehedbergella grigelisae Banner and sensis s.s. in having perforation cones and a shal-
Desai (sic), p. 154, pI. 2, figs 11-13 lower umbilicus. The presence of perforation
(?8-1O). cones in this subspecies is of particular interest
since these have not been previously recorded for
any other member of this genus.
Remarks: P. tuschepsensis grigelisi has five
chambers in the first whorl, reducing from four
and a half to four in the final whorl. The final Range: Early Barremian-Late Aptian. This
chambers are very slightly depressed (h/L = 0.92 subspecies was originally described from the
approx.). The umbilicus is small (u/uh = 0.19 Early Barremian to Late Aptian of the central
approx.) The aperture is a narrow slit, extending North Sea.
 Systematics 107

Praehedbergella handousi (Salaj, 1984) aperture is a high arch. Perforation cones are
visible on chambers of the last whorl.
(Plate 7.2, Figs 1-3; Figure 7.1)
1984 Caucasella handousi Salaj, p. 592, Figs Range: Late Barremian. P. pseudosigali was
2a--c. first described from the Late Barremian of the
central North Sea.
Remarks: Although the three figures assigned Praehedbergella ruka Banner, Copestake and
by Salaj (1984) to the holotype of C. handousi White, 1993 sensu stricto
seem to belong to different specimens (the dorsal
(Plate 7.2, Figs 7-9; Figure 7.1)
view not corresponding to the ventral and side
views), they should all be included under 1993 Praehedbergella ruka s.s. Banner,
Praehedbergella. Copestake and White, p. 6, pI. I, figs 2a--c.
P. handousi has four chambers in the last whorl.
The chambers of the last whorl enlarge rapidly Remarks: P. ruka s.s has five chambers in
and are only slightly depressed (hiL = 0.81 the penultimate whorl, but only four abutting the
approx.). The dorsal surface is fiat, the test being umbilicus in the final whorl. The dorsal surface is
ventro-dorsally compressed (dlt = 0.11 approx.). strongly convex (dlt =0.5 approx.), the chambers
The intra-extraumbilical aperture is a low slit, are slightly depressed (hiL = 0.8 approx.) and
possessing a thin rim. The umbilicus is broad and increase in size gradually with growth (bla = 1.8
shallow (uluh = 0.47 approx.). approx.). The umbilicus is small (uhlu = 0.15
This species differs from P. tuschepsensis approx.) and the aperture is a narrow slit.
(Antonova) in having a more compressed test and
a shallower umbilicus. Range: Early Aptian. This species was first
described from the Early Aptian of the central
North Sea.
Range: Early Hauterivian. P. handousi was first
described from the Early Hauterivian of Tunisia Praehedbergella ruka contritus Banner,
as Caucasella handousi. Banner et al., 1993, Copes take and White, 1993
referred to specimens from the Barremian of the (Plate 7.2 Figs 10-12; Figure 7.1)
central North Sea, resembling P. handousi but
smaller, as P. sp. cf. P. handousi. The identity of 1978 Hedbergella infracretacea gargasiana
these specimens could not be verified as their (Moullade), Caron, pp. 652-656, 658, pI.
initial chambers are not clearly seen. I, figs 7-9 (non Globigerina infracretacea
Glaessner gargasiana Moullade, 1961).
Praehedbergella pseudosigali Banner, Copestake 1982 ?Hedbergella hoterivica (Subbotina),
and White, 1993 Crittenden, pp. 30, 31, pI. 2, figs 1,2.
(Plate 7.2, Figs 4--6; Figure 7.1) 1993 Praehedbergella ruka contritus Banner,
Copestake and White, p. 6, pI. 1, figs 3a--c,
1992 Praehedbergella pseudosigali Banner, 4a, b.
Copestake and White, p. 6, pI. 2, figs la--c.
Remarks: P. ruka contritus is distinguished by
Remarks: P. pseudosigali has six and a half its strong dorsal convexity, depressed chambers
chambers in the penultimate whorl and four cham- (hiL = 0.8 approx.) in the last whorl, and a smooth
bers abutting the umbilicus in the last whorl. The test surface lacking perforation cones.
dorsal surface is slightly convex. The chambers Caron (1978) misidentified her material as H.
are symmetrical and increase in size gradually infracretacea gargasiana, a form which has five
with growth (bla = 2 approx.), those in the last and a half undepressed chambers in the final whorl
whorl become depressed (hiL = 0.85 approx.). The and the muricate test of Hedbergella; it is clearly
umbilicus is small (uluh = 0.29 approx.) and the conspecific with P. ruka contritus.
108 The flowering of the PraehedbergeUidae

A similar fonn has been figured by Crittenden Remarks: In 1966, Moullade described this
(1982) as ?H. hoterivica, but that species belongs species from the Hauterivian of La Drome and Les
to the genus Favusella (see also Banner and Desai, Basses-Alpes as well as from the Barremian of the
1988). Hautes-Alpes and (rarely) in the Lower Aptian.
His six accompanying photographs (1966, plate 7,
Range: Aptian. P. ruka contritus was first figs 20, 25) varied morphologically and would
described from the Early and Late Aptian of the now be considered to belong to different species.
central North Sea. Caron's specimens were from Fortunately, the holotype (1966, plate 7, figs
beds she believed to be Late Aptian and which 24-25; refigured on our Plate 7.3, Figs 7, 8) was
were in the lowest core obtained from DSDP Site photographed both ventrally and dorsally and
364, Angola Basin. Crittenden's specimens were shows the presence of subglobular chambers (hiL
from the Atherfield Clay (Early Aptian) of the Isle =0.6), separated by deep and narrow peripheral
of Wight, southern England. intercameral sutures (approaching 90), and a very
low, slit-like aperture. These characters can be
Praehedbergella ruka papillata Banner, used to recognize this species, sensu stricto; 'topo-
Copestake and White, 1993 type' specimens would be no use, because they
(Plate 7.3, Figs 1-3; Figure 7.1) could belong to any other species which happened
to occur in the same stratum at the same locality
1993 Praehedbergella ruka papillata Banner, as the holotype.
Copestake and White, p. 7, pI. I, figs 5a--c. The specimens figured by Moullade (1966, plate
7, figs 22, 23; Plate 7.4, Figs 3, 4, herein) as
Remarks: This species is identical to the typical paratypes were distinguished by Banner et al.
fonn of the species but has perforation cones (as (1993, p. 8) as P. sigali rasilis because this fonn
the name suggests). The aperture possesses a thick, has depressed chambers in the last whorl, which are
broad, heavy porticus. about half as high as long, moderately depressed
intercameral sutures, which fonn intercameral
Range: Early Aptian. P. ruka papillata was first angles of about 140 at the periphery of the test,
described from the Early Aptian of the central and a moderately lobate test outline (compared to
North Sea. the strongly lobate test of P. sigali s.s.). The holo-
type of P. sigali rasilis was chosen to be the
Praehedbergella sigali (Moullade, 1966) sensu specimen figured by Gorbachik (1986, plate 24, fig.
stricto I, 2; reproduced in our Plate 7.4, Figs I, 2) which
had been obtained by her from the Late Barremian
(plate 7.3, Figs 4-8, Plate 7.4, Figs 1-4; Figure
of the southwest Crimea. However, as we have
7.1)
not been able to obtain more specimens of this
1966 Hedbergella (Hedbergella) sigali Moull- fonn, we have not been able to add further to the
ade (pars), pp. 87, 88, pI. 7, figs 21-25 original diagnosis (1993) and for the moment
(non fig. 20). consider it a possible synonym of P. sigali.
1985 Hedbergella sigali Moullade; Caron, p. 59,
Fig. 25.21-22. Range: Early Barremian-Early Aptian. The
1986 Clavihedbergella sigali (Moullade), holotype of P. sigali s.s. was obtained by Sigal
Gorbachik, pI. 24, figs I, 2. (1966) from the Lower Barremian of the Hautes-
1993 Praehedbergella sigali (Moullade), Banner Alpes, southeastern France. The species has also
et al., p. 7, pI. 2, figs 2a--c. been shown to range from the Early Barremian to
?1993 Praehedbergella sigali (Moullade) rasilis the Early Aptian in the central North Sea area
Banner, Copestake and White, p. 8. (Banner et al., 1993, p. 8). BouDagher-Fadel
1995 Praehedbergella sigali (Moullade), (1995) obtained it from the Early Barremian of
BouDagher-Fadel, p. 142, pI. 2, figs 1,2. Tunisia.
 Systematics 109

Praehedbergella sigali (Moullade) compacta Yorkshire, England (Schackoina cabri Zone),

Banner, Copestake and White, 1993 from Japan and the Gargasian stratotype (Late
Aptian, Globigerinelloides ferreolensis Zone),
(Plate 7.3, Figs 9-12; Figure 7.1) Vaucluse, SE France.
1966 Hedbergella (Hedbergella) sigali Moullade Praehedbergella yakovlevae sp .nov.
(pars), pp. 87, 88, pI. 7, fig. 20 only.
1993 Praehedbergella sigali (Moullade) com- (Plate 7.4, Figs 8-10; Figure 7.1)
pacta Banner et al., p. 8, pI. 2, fig. 3a-c.
1993 Praehedbergella tuschepsensis (Antonova),
Banner, Copestake and White, pp. 8, 9,
Remarks: This subspecies differs from the
pI. 2, figs 4a-c (non Globigerina tuschep-
typical form of the species in having peripheral
sensis Antonova, 1964).
intercameral sutural angles of about 150 and more
0

depressed chambers in the last whorl (hlL:::; 0.8

Diagnosis: Praehedbergella with four and a half
approx.), producing a more compact test. The
chambers in the last whorl but with only four
chambers of the last whorl become less depressed
surrounding the umbilicus; the dorsal surface is
during growth, and the final chamber is only
flat but the dorsal intercameral sutures are oblique
slightly lower than long.
to the spiral suture (not radially arranged) and the
A specimen of H. (H.) sigali figured by
dorsal chambers are depressed (hL:::; 0.6 approx.)
Moullade (1966, plate 7, fig. 20), in spite of a
and highest posteriorly (1/L2 :::; 2.6 approx.). The
rather poor optical photograph taken at a slightly
umbilicus is broad (u/uh:::; 0.4 approx.) The slit-
oblique angle (our Plate 7.3, Fig. 9), resembles the
subspecies enough to be included here. like aperture possesses a broad porticus. The
chambers of the last whorl are quite thick (hit:::;
0.6 approx.). No perforation cones are yet known.
Range: Known from the Early Barremian to the
Early Aptian of the central North Sea area.
Provenance of the holotype: The holotype was
Moullade's (1966) specimen came from the Early
obtained from central North Sea Well 16/28-6RE,
Barremian of SE France.
4150.5 m subsurface, Early Aptian. Deposited in
Praehedbergella tatianae Banner and Desai, The Natural History Museum, London, no. P 52733.
1988
Remarks: This is the only species of Praehed-
(Plate 7.4, Figs 5-7; Figure 7.1)
bergella in which the chambers of the last whorl
1974 Hedbergella sp. cf H. sigali Moullade; are at their highest point posteriorly. Because of
Longoria, pI. 22, figs 1-5. this character, specimens from the North Sea Wells
1982 Hedbergella cf H. sigali Moullade; Obata were originally called P. tuschepsensis (Antonova)
et al., p. 156, pI. 4, figs 2a-c. as Gorbachik's figure (1986, plate 25, fig. 1), called
1988 Praehedbergella tatianae Banner and by her by that species name, also had posteriorly
Desai, p. 152, pI. 3, figs 1-3. high chambers. However, examination of type
material of P. tuschepsensis, as noted above in the
Remarks: This species is characterized by its Introduction (p. 105), has shown that the specimen
very depressed chambers (h/L:::; 0.4 approx.) and figured by Gorbachik is really a Blefuscuiana. This
very broad peripheral intercameral sutural angles Praehedbergella therefore needs a new name. We
(160 0 approx.). No specimens with perforation are pleased to dedicate it to Dr Svetlana Yakovleva
cones are yet known. The umbilicus is small and of St Petersburg.
the slit-like aperture possesses a broad porticus.
Range: P. yakovlevae occurs from the Early
Range: Late Aptian. This species is known only Barremian into the Early Aptian of the central
from the early Late Aptian of Speeton, North North Sea area.
110 The flowering of the Praehedbergellidae

7.4 CONCLUDING COMMENTS appeared abutting the umbilicus, and these are
now referred to the genus Blefuscuiana. The four-
Praehedbergella species range in age from Early chambered Praehedbergella died out in the
Hauterivian to Late Aptian, but the high spired Aptian, but gave rise to similarly four-chambered
forms (P. ruka s.l.) are as yet not known before but planispiral Blowiella, the first Schackoinidae,
the Early Aptian. In the Barremian, five chambers in the late Early Barremian (Figure 5.2).
Plate 7.1

Figures 1-7 Praehedbergella tuschepsensis (Antonova, 1964) sensu stricto emend. 1-3 Early Barremian, along the Tuscheps
River, northwestern Caucasus, holotype (from Antonova, 1964, pI. 12, figs 3a-{;), deposited in VNIGRI, St
Petersburg, Russia, no. 6227: ventral view showing clearly four chambers abutting the umbilicus, the dorsal
and peripheral views (x240); 4-7, same holotype figured here by SEM for the first time: 4-6, ventral view
showing a broad umbilicus, the flat dorsal side and the peripheral view (x300); 7, enlargement of the first whorls
of the dorsal side showing a smooth microperforate test (x 1000).
Figures 8-11 Praehedbergella tuschepsensis (Antonova) perforare Banner, Copestake and White, 1993. Early Aptian, Well
20/2-2, 8300 ft, central North Sea; holotype deposited in The Natural History Museum, London, no. PF 52758:
8-10, ventral, peripheral and dorsal views (x240); 11, enlargement of the surface to show perforation cones
(x2500).
Figures 12-14 Praehedbergella tuschepsensis (Antonova) grigelisi (Banner and Desai, 1988), earliest Late Aptian, Schackoina
cabri Zone, Speeton Cliff, NE England, deposited in The Natural History Museum, London, nos PF 52129
(holotype), PF 52115 (paratype): 12, 14, ventral view of holotype showing a narrow umbilicus (x270), 14,
enlargement showing the narrow slit aperture furnished with a porticus (x540); 13, peripheral view of paratype
showing a convex dorsal side (x340).
Plate 7.2

Figures 1-3 Praehedbergella handousi (Salaj, 1984). Early Hauterivian, Jebel Oust, Tunisia, holotype deposited in the collec-
tions of the Dionys St6r Institute of Geology, Bratislava, no. T-IO (from Salaj, 1984, Figs 2a-c): dorsal, periph-
eral and ventral views (x165 approx).
Figures 4-6 Praehedbergella pseudosigali Banner, Copestake and White, 1993. Late Barremian, Well 2012-2, 8370 ft, central
North Sea, holotype deposited in The Natural History Museum, London, no. PF 52735: ventral, peripheral and
dorsal views (x295).
Figures 7-9 Praehedbergella ruka Banner, Copestake and White, 1993 sensu stricto. Early Aptian, Well 16/28-6RE, 4157.5
m, central North Sea, holotype deposited in The Natural History Museum, no. PF 52739: dorsal, peripheral and
ventral views (x315).
Figures 10-12 Praehedbergella ruka contritus Banner, Copestake and White, 1993. Late Aptian, Well 16/28-6RE, 3989.5 m,
central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52737: dorsal. periph-
eral and ventral views (x355).
Plate 7.3

Figures 1'--3 Praehedbergella ruka papillata Banner, Copestake and White, 1993. Early Aptian, Well 16/28-6RE, 4150 m,
central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52736: dorsal, periph-
eral and ventral views (x290).
Figures 4-8 Praehedbergella sigali (Moullade, 1966) sensu stricto. 4-6, Early Barremian, Well 20/2-2, 8680 ft, central North
Sea, holotype deposited in The Natural History Museum, London, no. PF 52742: ventral, dorsal and peripheral
views (x325). 7, 8, Early Barremian, Hautes-Alpes, SE France, holotype deposited in the Department of Earth
Sciences, University of Lyon, France, no. 50 113 (from Moullade, 1966, pI. 7, figs 24, 25): ventral and dorsal
views (x 100).
Figures 9-12 Praehedbergella sigali (Moullade) compacta Banner, Copestake and White, 1993. 9, Early Barremian, Hautes-
Alpes, SE France, paratype of P. sigali (from Moullade, 1966, pI. 7, fig. 20), deposited in the Department of
Earth Sciences, University of Lyon, France, no. 50 114: ventral side (xlOO). 10-12, Early Barremian, Well 20/2-
2,8680 ft, central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52746: dorsal,
ventral and peripheral views (x365);
Plate 7.4

Figures 1-4 Praehedbergella sigali (Moullade, 1966) sensu stricto, 1,2, Late Barremian, Kacha River, southwestern Crimea,
Russia (from Gorbachik, 1986, pI. 24, figs 1,2), deposited in the Geology Department of Moscow State University,
Russia, nos 78, 79: dorsal and ventral views (x400). 3, 4, Early Barremian, Hautes-Alpes, SE France, paratypes
deposited in the Department of Earth Sciences, University of Lyon, France, nos 50 116, 50 117 (from Moullade,
1966, pI. 7, figs 22, 23): ventral views (x 100).
Figures 5-7 Praehedbergella tatianae Banner and Desai, 1988. Early Late Aptian, Late Schackoina cabri Zone, Speeton Cliff,
NE England, holotype deposited in The Natural History Museum, London, no. PF 52131: 5,6, ventral and dorsal
views (x270), 7, surface detail with microperforations (xI200).
Figures 8-10 Praehedbergella yakovlevae sp.nov. early Aptian, Well 16/28-6RE, 4150.5 m, central North Sea, holotype
deposited in The Natural History Museum, London, no. PF 52733: ventral, peripheral and dorsal views (x240).
 8
Blefuscuiana, the longest
ranging and most diverse
of the Praehedbergellidae
M.K. BouDagher-Fadel, F.T. Banner
and J .E. Whittaker

8.1 INTRODUCTION Species of Blefuscuiana are normally much

smaller than those of Hedbergella (just as
In the Early Barremian, the trochospiral taxon Tenuitella is characteristically smaller than
Praehedbergella, with four chambers abutting the Paragloborotalia in the Tertiary). The microper-
umbilicus, gave rise to Blefuscuiana by increasing forations of Blefuscuiana (Plate 1.1, Fig. 2) are
the number of chambers in the whorl (Figure 5.2). characteristically much smaller than the macro-
By Aptian time, a diversity of many-chambered perforations of Hedbergella (Plate 1.1, Fig. 7), and
species of Blefuscuiana was well established (Figure much more irregularly distributed on any part of
8.1), and the genus lasted well into the Early Danian. the test surface. The total absence of muricae
Blefuscuiana is trochospiral, smooth and over the test surface of Blefuscuiana means that
microperforate (but sometimes with perforation muricocarinae could never develop. Therefore,
cones). The perforation cones are hollow as they although Praeglobotruncana (Plate ll.l, Figs
are developed over the microperforations (e.g. 5-10) could evolve from Hedbergella (Plate 11.1,
Blefuscuiana infracretacea (Glaessner) s.s, Plate Figs 1-4), Blefuscuiana was unable to follow an
8.7, Figs 1-8); this structure is not known again analogous evolution. No imperforate peripheral
in the history of the Globigerinina until they re- band is known in any species of Blefuscuiana,
appeared in the heterohelicids: Guembilitria of the although imperforate areas could develop (appar-
Maastrichtian (Smith and Pessagno, 1973, plate 1, ently sporadically) around the umbilical margins
figs 1, 3, 5, 8), and Cassigerinella (as described of some species (a character not yet known in
by Li, 1986) of the Oligocene. Hedbergella or Praeglobotruncana). Evolution of
In the Late Aptian, the acquisition of macro- gross morphology, where it was unaffected by the
perforations and test muricae, as shown by lack of muricae, occurred along lines analogous to
BouDagher-Fadel et al., 1997a, led to the evolu- those of the Hedbergellidae, resulting in gross
tion of Hedbergella. homeomorphs between the two families. In the
122 Blefuscuiana of the Praehedbergellidae

127 121 112.2 Ma

/
BARREMIAN APTIAN ALBIAN
EARLY LATE EARLY LATE EARLY LATE

Blefusculana alblana
B. aptlana s.s.
B. aptlana orlentalls
B. convexa
B. damlnlae
B. excelsa s.s.
B. excelsa cumulus
B. gorbachlkae
B. hexacamerata
B. hlspanlae
B. Infracretacea s.s.

-- B. Infr. occidentails
B. kuznetsovae
B. laculata s.s.
B. laculata alobata
B.mltra
B. multlcamerata
B. occulta s.s.
B. occ. perforocculta
B. praetrocoldea
B.prlmare
B. rudls s.s.
B. rudls pararudls
B. speetonensls s.s.
B. speet. tunlslensls
B.sp.

Figure 8.1 Stratigraphical distribution of Early Cretaceous Blefuscuiana. Chronostratigraphy and Absolute Time Scale after
Gradstein et al. (1995).
 Key to the species of Blefuscuiana 123

latest Barremian Blefuscuiana species gave rise to 1.2.2.1.1 Chambers of the last whorl slowly
Lilliputianella (homeomorph of Clavihedbergella) enlarging with small umbilicus
by radially elongating their chambers (Figure 5.2), 1.2.2.1.1.1 Test less lobate, ventral chamber
and in the Late Aptian developed streptospirality sides not umbilically extended
(Wondersella, Plate 9.4), uniquely in the Mesozoic. B. infracretacea s.s.
Although the morphological extremes of 1.2.2.1.1.2 Test more lobate, ventral chamber
Blefuscuiana (e.g. B. excelsa (Longoria) cumulus sides umbilically extended
Banner, Copestake and White, Plate 8.5, Figs 7-12 B. gorbachikae
and B. multicamerata Banner and Desai, Plate 8.8, 1.2.2.1.2 Chambers of the last whorl rapidly
Figs 9, 10) are readily separable, there is such a enlarging with larger umbilicus
wide diversity of other forms that the casual view B. albiana
may give the impression that all are 'varieties' of 1.2.2.2 Chambers asymmetrical, highest pos-
one (or at best a few) species. The taxonomy we teriorly B. sp. (Plate 8.12,
have employed is designed to enable the recording Figs 7-9)
of all these different forms, so that their distribu- 1.2.3 Last chambers not depressed but as high
tion and stratigraphic range may also be docu- as long, tending to be subglobular
mented. In anyone sample, two or more different 1.2.3.1 Dorsal side flat to concave, peripheral
taxa may be found to intergrade but stratigraphi- intercameral sutural angles narrow
cally younger samples may well show that only B. mitra
one of these taxa persists. So, unless precise taxo- 1.2.3.2 Dorsal side weakly convex, peripheral
nomic records are maintained, the diversity intercameral sutural angles relatively broad
changes will not be recorded either, and biostrati- 1.2.3.2.1 Test thick, umbilicus small
graphic and evolutionary information will be lost. B. daminiae (= B. whittaken)
Some of the new taxa may be regarded by the 1.2.3.2.2 Test weakly compressed, umbilicus
reader as synonyms, but it must be remembered broad
that new data from new assemblages may show 1.2.3.2.2.1 Periphery more lobate, peripheral
that their distinction would be valid. intercameral sutural angles relatively narrow
B. laculata s.s. (= B. praesimilis)
1.2.3.2.2.2 Periphery less lobate, test more
8.2 KEY TO THE SPECIES OF quadrate, intercameral sutural angles relatively
BLEFUSCUIANA broad
B. laculata alobata
1 Five chambers abut the umbilicus and five 2 Six chambers abutting the umbilicus and six
chambers are visible dorsally in the last whorl to seven are visible dorsally in the last whorl
1.1 Dorsal side strongly convex, spire very 2.1 Chambers very depressed, peripheral
high sutural intercameral angles broad
B. excelsa cumulus 2.1.1 Dorsal side convex, umbilicus small
1.2 Dorsal side flat or weakly convex B. excelsa s.s.
1.2.1 Chambers very depressed, peripheral 2.1.2 Dorsal side flat, umbilicus broad
sutural angles very broad B. hispaniae
1.2.1.1 Test compressed 2.2 Last chambers slightly depressed, periph-
=
B. convexa B. moulladei eral sutural intercameral angles less broad
1.2.1.2 Test thick, not compressed 2.2.1 Test very thick, umbilicus small
B. rudis s.s. B. rudis pararudis
1.2.2 Chambers weakly depressed, peripheral 2.2.2 Test less thick, umbilicus broader
sutural angles less broad 2.2.2.1 Dorsal side flat to concave, umbilicus
1.2.2.1 Chambers dorsally symmetrical, very broad
highest in mid-point B. primare
124 Blefuscuiana of the Praehedbergellidae

2.2.2.2 Dorsal side weakly convex, umbilicus 8.3 SYSTEMATICS

less broad
2.2.2.2.1 Dorsal chamber sides less long, Superfamily GLOBIGERINACEA Carpenter,
seven chambers visible dorsally in last whorl; Parker and Jones, 1862
aperture extends fully to periphery of penulti-
mate whorl, with a broad portical lip Family PRAEHEDBERGELLIDAE Banner
B. praetrocoidea and Desai, 1988
2.2.2.2.2 Dorsal chamber sides longer, six
Genus Blefuscuiana Banner and Desai, 1988
visible dorsally in the last whorl; aperture
nearly reaches periphery with narrow portical Type species Blefuscuiana kuznetsovae Banner
lip and Desai, 1988
B. infracretacea occidentalis
Blefuscuiana kuznetsovae Banner and Desai,
2.3 Chambers on dorsal side virtually as high
1988
as long, little if at all depressed
2.3.1 Dorsal side flat to concave (Plate 8.1, Figs 1-5; Figure 8.1)
2.3.1.1 Ventral chambers imperforate around
1988 Blefuscuiana kuznetsovae Banner and
the umbilicus; aperture extends to periphery of
Desai, pp. 156-158, pI. 3, fig. 10; pI. 4,
penultimate whorl
figs 1, 2, 4 (not fig. 3).
B. speetonensis s.s.
2.3.1.2 Aperture extends beyond periphery of
Remarks: B. kuznetsovae has seven chambers
penultimate whorl to reach the dorsal side
abutting the umbilicus in the last whorl. Last
B. speetonensis tunisiensis
chambers are inflated, subglobular (h/L = 1
2.3.2 Dorsal side flat to slightly convex
approx.), and almost as high as long (L1 = 2). The
2.3.2.1 On dorsal side, chambers enlarge more
umbilicus is broad (u/uh = 0.27 approx.) and
rapidly, early whorls occupy about one quarter
shallow. The aperture is a low arch, from the
of dorsal surface
umbilicus to the periphery of the penultimate
2.3.2.1.1 Test thicker, umbilicus smaller
whorl, with a distinct porticus which broadens
B. hexacamerata
slightly posteriorly; relict apertures and their
2.3.2.1.2 Test thinner, umbilicus broader
portici are open for at least half of the last
B. aptiana s.l.
whorl.
2.3.2.1.2.1 No perforation cones
B. aptiana s.s. = (B. occulta
Range: Late Aptian. This species was first
quinquecamerata)
described from the Late Aptian of the Speeton
2.3.2.1.2.2 With perforation cones
Cliffs, Filey Bay, North Yorkshire, England.
B. aptiana orientalis
2.3.2.2 On dorsal side, chambers enlarge more Blefuscuiana albiana BouDagher-Fadel et aZ.,
slowly, early whorls occupy half of dorsal side 1996
B. occulta s.l.
(Plate 8.1, figs 6-9; Figures 8.1, 8.3)
2.3.2.2.1 With no perforation cones
B. occulta s.s. 1969 Hedbergella infracretacea (Glaessner),
2.3.2.2.2 With perforation cones Gorbachik and Kretchmar, pp. 50, 51,
B. occulta perforocculta pI. 1, figs 5a-c (non Globigerina infracre-
3 With seven subglobular chambers in final whorl tacea Glaessner, 1937).
B. kuznetsovae 1975 Hedbergella infracretacea (Glaessner),
4 With eight subglobular chambers in final whorl Magniez-Jannin (pars), p. 250, pI. 17, figs
B. multicamerata 4,6, 12, 15.
1986 Hedbergella injracretacea (Glaessner),
Gorbachik, p. 235, pI. 19, figs 1,2.
 Systematics 125

1996 Blefuscuiana albiana BouDagher-Fadel 1988 Blefuscuiana aptiana (Bartenstein), Banner

et al., p. 253, Fig. 4(6-9). and Desai, p. 158, pI. 5, figs 1-3.
1988 Blefuscuiana occulta (Longoria) quinque-
Remarks: B. albiana has five subglobular (hiL = camerata Banner and Desai, p. 162, pI. 7,
0.8 approx.) chambers in the last whorl, which figs 9-11.
enlarge rapidly with growth (bla = 2 approx.), so 1990 Hedbergella infracretacea (Glaessner),
that the addition of a new whorl increases the dor- Weidich (pars), p. 166, pI. 56, figs 19-21
sal test diameter by almost twice. The umbilicus is (not 24-26).
small, as in B. infracretacea (uluh = 0.3 approx.) 1993 Blefuscuiana aptiana (Bartenstein),
and is about 18 to 20% of the ventral test diame- Banner, Copestake and White, p. 9, figs
ter. The aperture is slit-like with a thin porticus. The 4a-b, 5, 6a-b.
wall is smooth; the microperforations are irregu- 1996 Blefuscuiana aptiana (Bartenstein) s.s.,
larly spaced and are about 2 /-Lm in diameter (twice BouDagher-Fadel et al., p. 252, Fig. 4
as large as those of B. aptiana (Bartenstein) s.l. but (1-5).
about half the size as those of true Hedbergella).
There appears to be a slight but progressive Remarks: B. aptiana s.s. has six subglobular
increase in test size from B. aptiana (Bartenstein) chambers (hiL = 1 approx.) in the final whorl.
s.s. (190-270/-Lm) and B. aptiana (Bartenstein) The umbilicus is broad, the aperture is a narrow
orientalis BouDagher-Fadel et al. (210-270 /-Lm) in intraumbilical-extraumbilica1 slit bordered by a
the Aptian to B. albiana (220-290 /-Lm) in the porticus which is absent in B. aptiana
Albian. It has already been suggested that B. (Bartenstein) orientalis BouDagher-Fadel et al.
albiana evolved from B. aptiana S.s. above the base
of the Albian (BouDagher-Fadel et al., 1996).
Range: Early Barremian-Early Albian. B.
aptiana s.s. ranges from the Early Barremian
Range: Latest Early Albian-Late Albian. The
to the Late Aptian in the North Sea and it
holotype and paratypes of B. albiana were
is recorded from the Early Aptian to Early Albian
obtained from the Upper Albian (AI 3) of Pre-
in north-central Europe (Bartenstein, 1965). We
doroshnaya, SW Crimea. Similar specimens were
have also identified specimens sent to us by
recorded by Magniez-Jannin (1995) from the
Dr M. Bubik from the Early Barremian-Aptian
'Middle Albian' (probably latest part of the Early
of the West Carpathians, Outer Flysch Belt,
Albian) of the Aube, France.
Magura group of nappes, Hluk Formation, as
Blefuscuiana aptiana (Bartenstein, 1965) sensu belonging to B. aptiana s.s. Similar specimens
stricto were figured by Weidich (1990) from the Late
Aptian of Bavaria.
(Plate 1.1, Fig. 2, Plate 8.2, Figs 1-8; Figures
8.1,8.3) Blefuscuiana aptiana (Bartenstein) orientalis
1938 ?Globigerina sp. D9 Hecht, p. 17, pI. 23, BouDagher-Fadel et al., 1996
fig. 60. (Plate 1.1, Figs 3, 4, Plate 8.2, Figs 9-11, Plate
1938 Globigerina sp. D9 Hecht, p. 17, pI. 23, 8.3, Figs 1-8; Figures 8.1,8.3)
figs 61-63.
1965 Hedbergella aptiana Bartenstein, pp. 347, 1986 Hedbergella aptica (Agalarova), Gorba-
348, text-figs 3-6. chik, pp. 94, 234, pI. 14, figs 4, 5 (not figs
1975 Hedbergella infracretacea (Glaessner), 2,3).
Magniez-Jannin (pars), pp. 250-254, pI. 1993 Blefuscuiana aptiana (Agalarova) implana
17, figs 1-3,9, 10, 18. Banner, Copestake and White (pars), p. 10,
1979 Hedbergella aptiana (Bartenstein), Sigal, pI. 7, fig. 2 (paratype); (not fig. 3, holo-
p. 318, pI. 2, figs 24, 25. type).
126 Blefuscuiana of the PraehedbergeUidae

1993 Blefuscuiana aptica (Agalarova), Shahin, B. convexa has five very depressed chambers
p. 421, pI. 5, fig. 3. (h/L =approx.) in the last whorl. The umbilicus is
1996 Blefuscuiana aptiana (Bartenstein) orien- narrow (uluh = 0.25 approx.). The aperture is a
talis BouDagher-Fadel et al., p. 252, Fig. narrow intraumbilical-extraumbilical slit.
2(4-12), Fig. 3(1-8).
Range: Early Late Aptian. Described by
Remarks: B. aptiana orientalis has the same Longoria (1974) from the Schackoina cabri Zone,
shape of test as B. aptiana (Bartenstein) s.s. with of the la Drome region of SE France. We have
a flat to slightly concave dorsal side, six subglob- also identified it (Plate 8.4, Figs 7, 8) in material
ular chambers in the final whorl (with the last of the same age sent to us by Dr M. Bubik from
chambers being as high as long, h/L = 1 approx.), the Hluck Formation, Outer Flysch Belt, Magura
but it has perforation cones on the early chambers group of nappes, West Carpathians.
of the last whorl and there is no porticus bordering Blefuscuiana daminiae Copestake and White,
the aperture. 1993

Range: Within the Late Barremian-Early Late (plate 8.5, Figs 1-3, Plate 8.12, Figs 4-6;
Aptian. B. aptiana orientalis was first described Figure 8.1)
from the Early Aptian of Oued Guetta in Tunisia. 1993 Blefuscuiana daminiae Banner, Copestake
It was figured by Shahin (1993) as B. aptica and White, p. 10, pI. 3, figs 6a-c; pI. 4,
(Agalarova) from the Early Aptian of Bardawil figs la, b; pI. 4, figs la, b, 3a-c.
area, northern Sinai in Egypt. We have found it 1993 Blefuscuiana whittakeri Banner, Copestake
in the Early Aptian (Ap2) of SW Crimea and White, p. 14, pI. 5, figs 3, 4.
(Krasnaya) and Azerbaijan and the Late Barremian 1997b Blefuscuiana daminiae Banner, Copestake
of the North Sea. and White; BouDagher-Fadel et al., pI. 1,
Blefuscuiana convexa (Longoria, 1974) figs 8, 9 (in press).

(Plate 8.4, Figs 1-8; Figure 8.1) Remarks: B. daminiae has five subglobular (h/L
1974 Loeblichella convexa Longoria, p. 73, pI. =0.9 approx.) chambers in the last whorl. The test
14, figs 12-14. is thick, the umbilicus is small (u/uh =0.5 approx.)
1974 Loeblichella moulladei Longoria, p. 73, pI. and the aperture is narrow.
14, figs 9-11, 15-17.
Range: Late Barremian to Early Aptian. This
species was first described from the North Sea area.
Remarks: Loeblich and Tappan (1988, p. 704)
It was recorded in the Kacha River, SW Crimea,
considered Loeblichella Pessagno (1967) as a
Azerbaijan and Tunisia (BouDagher-Fadel et al.,
genus of uncertain status; the types being prob-
1997b).
ably internal moulds and poorly preserved. Also,
Longoria's specimens lack the sutural apertures on Blefuscuiana excelsa (Longoria, 1974) sensu
the spiral side originally said to occur on the type stricto
species, Praeglobotruncana hessi hessi. According
(Plate 8.5, figs 4-6; Figure 8.1)
to Longoria (1974), his L. convexa differs from L.
moulladei only by having a less convex dorsal 1974 Hedbergella excelsa Longoria, pp. 55, 56,
side. A close study of the type specimens of the pI. 18, figs 6-11,14-16.
two species (reproduced in Plate 8.4, Figs 1-6)
shows that the degree of convexity of the dorsal Remarks: B. excelsa s.s. has six very depressed
side, in both species, is the same. We regard them chambers (h/L =0.2 approx.) abutting the
as synonyms and, as first revising authors, select umbilicus, a convex dorsal side (dlt =0.4 approx.)
convexa has having priority. and a small umbilicus (u/uh =0.25 approx.).
 Systematics 127

Range: Early Aptian. First described by 1993 Blefuscuiana gorbachikae (Longoria),

Longoria (1974) from the lower part of the la Shahin, p. 422, pI. 5, fig. 4.
Drome section, SE France. It is not as yet known 1995 Blefuscuiana gorbachikae (Longoria),
from anywhere else. BouDagher-Fadel, p. 143, pI. 2, fig. 5.
1997b Blefuscuiana gorbachikae (Longoria),
Blefuscuiana excelsa (Longoria) cumulus
BouDagher-Fadel et al., pI. 3, figs 4-6 (in
Banner, Copestake and White, 1993
press).
(Plate 8.5, Figs 7-12; Figure 8.1)
Remarks: This species is characterized by its
1993 Blefuscuiana excelsa (Longoria) cumulus
lobate test, later chambers not depressed (hiL ,; 1.2
Banner, Copestake and White, p. 11, pI. 6,
approx.) and strongly overlapping the umbilicus.
figs 1a-c, 2a-c.
1997b Blefuscuiana excelsa (Longoria) cumulus
Range: Early Aptian to Early Albian. B. gorba-
Banner, Copestake and White; BouDagher-
chikae was first described by Longoria from
Fadel et al., pI. 2, figs 1-8 (in press).
Mexico, southeastern France and Spain. It was
also recorded from the Late Aptian-latest Early
Remarks: This subspecies has five slightly
Albian of Bavaria (Weidich, 1990). We have
depressed (hiL = 0.6 approx.) chambers abutting
found it extensively in Tunisia (BouDagher-Fadel,
the umbilicus, a strongly convex dorsal side (dlt
1995), Azerbaijan and Crimea (BouDagher-
= 0.4 approx.) (this convexity being more
Fadel et al., 1997b) throughout its stratigraphic
pronounced in some specimens, Plate 8.5, Fig. 10),
range.
a small and deep umbilicus and a narrow aperture
with a thin rim. Blefuscuiana hexacamerata sp.nov.
(Plate 8.6, Figs 7, 8; Figure 8.1)
Range: Early Aptian. This species was des-
cribed from the Early Aptian of the Central North
Derivation of Name: After the number of cham-
Sea area. It was subsequently found in the Early
bers in the final adult whorl.
Aptian of the Kacha River and Azerbaijan
(BouDagher et al., 1997b). We have also identi-
Diagnosis: Characterized by its flat to concave
fied this species in the Early Aptian of the Hluck
spiral side, and by having six chambers in the final
Formation of the West Carpathians.
whorl.
Blefuscuiana gorbachikae (Longoria, 1974)
Holotype: PF 53079, deposited in The Natural
(Plate 8.6, Figs 1-6; Figure 8.1)
History Museum, London. From sample N120,
1974 Hedbergella gorbachikae Longoria, pp. Kacha River, SW Crimea, Late Aptian.
56-58, pI. 15, figs 1-16.
1985 Hedbergella gorbachikae Longoria, Caron, Description: Test about 230 j..Lm at its maximum
p. 31, 59, figs 25.8-9. diameter, comprising two to three whorls coiled
1986 Hedbergella gorbachikae Longoria, in a low trochospire, partially evolute and concave
Gorbachik, p. 94, pI. 16, figs 1, 2. on spiral side with six chambers in the last whorl,
1988 Blefuscuiana gorbachikae (Longoria), inflated, subglobular (hiL = 1 approx.). The first
Banner and Desai, p. 160, pI. 5, figs 8-12. chambers of the last whorl, when seen on the spiral
1990 Hedbergella gorbachikae Longoria, side, are slightly longer than high, but as high as
Weidich, p. 166, pI. 56, figs 13-15, pI. 58, long in the last three chambers. The dorsal side is
figs 1-3. flat to concave. The umbilicus is small (uluh = 0.3
1993 Blefuscuiana gorbachikae (Longoria), approx.) and deep and the aperture is a narrow
Banner, Copestake and White, p. 11, pI. 6, slit. Perforation cones have not been found on this
figs 3a-c. species.
128 Blefuscuiana of the Praehedbergellidae

Remarks: B. hexacamerata differs from B. 1966 Hedbergella infracretacea (Glaessner),

infracretacea (Glaessner) occidentalis BouDagher- Glaessner, pp. 179-184, pI. 1, figs la-3c.
Fadel et al. by its partially evolute and concave 1969 Hedbergella aptica (Agalarova), Gorba-
spiral side and by its higher number of chambers chik and Kretchmar, pp. 50, 51, pI. 1, figs
on the umbilical side, and from B. gorbachikae la-c.
(Longoria) by the higher number of chambers in 1986 Hedbergella aptica (Agalarova), Gorba-
the last whorl and by the absence of an overhanging chik, p. 94, pI. 14, figs 4, 5 (not 2, 3).
last chamber. The test is thicker, and its umbilicus 1988 Blefuscuiana cf. aptica (Agalarova), Banner
is smaller than in B. aptiana (Bartenstein). and Desai, p. 160, pI. 3, figs 4, Sa, b.
1993 Blefuscuiana infracretacea (Glaessner)
Range: Late Aptian. So far only known from the sensu stricto Banner, Copestake and
type locality, Kacha River, SW Crimea. White, p. 11, pI. 4, fig. 2, pI. 8, fig. 1.
1995 Hedbergella delrioensis (Carsey), Weiss,
Blefuscuiana hispaniae (Longoria, 1974)
pI. 1, figs 1, 2.
(Plate 8.6, Figs 9-11; Figures 8.1,8.3) 1995 Hedbergella infracretacea (Glaessner),
Weiss, pI. 1, fig. 4.
1971 Hedbergella infracretacea (Glaessner),
1996 Blefuscuiana infracretacea (Glaessner),
Kuhry, p. 230, pI. 2, figs 3a-c.
BouDagher-Fadel et al., pp. 246-249, fig.
1974 Hedbergella hispaniae, Longoria, p. 58, pI.
1(1-8), fig. 2(1-3).
19, figs 4--8, 19-20.
1997b Blefuscuiana infracretacea s.s (Glaessner),
BouDagher-Fadel et al., pI. 3, fig. 1 (in
Remarks: This species has six very depressed
press).
chambers (hiL = 0.5 approx.) abutting the
umbilicus, a flat dorsal side and a broad umbilicus
Remarks: B. infracretacea s.s. has five cham-
(uluh = 0.4 approx.)
bers in the last whorl and strongly developed
perforation cones. It grades into infracretacea
Range: Late Aptian. Longoria recorded this
(Glaessner) occidentalis BouDagher-Fadel et al.
species from the la Perra Formation at La Boca
which has five chambers in the last whorl and
Canyon and at la Dr6me, SE France. Kuhry
lacks perforation cones. The existence of this
recorded the occurrence of similar forms from the
morphological grade is exemplified by the speci-
Argos Formation of Spain, as H. infracretacea
men shown on Plate 8.7, Figs 12-15, which still
(Glaessner). We have identified B. hispaniae from
has five chambers in the whorl but which lacks
specimens collected by Dr M. Bubik from the Late
perforation cones except for their weak develop-
Aptian of the Hluk Formation of the West
ment on the first chamber of the last whorl. This
Carpathians.
transitional form demonstrates one of the reasons
Blefuscuiana infracretacea (Glaessner, 1937) why we consider the presence or absence of per-
sensu stricto foration cones to be indicative of sub specific
distinction only. Weiss (1995) illustrated a similar
(Plate 8.7, Figs 1-8; Figures 8.1, 8.3)
form (as Hedbergella delrioensis) from the Early
1937 Globigerina infracretacea Glaessner, p. to Middle Aptian of the Wiechendorf 1/86
28, text-fig. 1. Borehole, north of Hannover.
1951 Globigerina aptica Agalarova, p. 49, pI. 8,
figs 9-11. Range: Early Aptian (not earliest part)-Early
1960 Globigerina infracretacea Glaessner gar- Albian. This species is known from the Aptian of
gasiana Moullade, p. 136, pI. 2, figs 21, Azerbaijan, Crimea, SE France, Germany and the
23-25. (probably Early) Albian of the Caucasus. It is not
1960 Globigerina infracretacea Glaessner gar- known from further west (e.g. the North Sea area
gasiana Moullade, p. 214. and Tunisia) where the specimens of identical
 Systematics 129

chamber form lack perforation cones (B. infra- by Dr M. Bubick in the Early Aptian of the Hluk
cretacea occidentalis, see BouDagher-Fadel et al., Formation, West Carpathians, as belonging to this
1996). subspecies.
Blefuscuiana infracretacea (Glaessner) occiden- Blefuscuiana laculata Banner, Copestake and
talis BouDagher-Fade1 et al., 1996 White, 1993 sensu stricto

(Plate 8.7, Figs 9-15; Figures 8.1-8.3) (Plate 8.8, Figs 1-3, Plate 8.9, Figs 1-3; Figure
8.1)
1980 Hedbergella aptica (Agalarova), Gorba-
chik, pI. 3, figs 2, 3. (non Globigerina 1979 Clavihedbergella eocretacea Neagu; Sigal,
aptica Agalarova, 1951 == B. infracretacea p. 290, pI. 3, figs 31 (?30).
s.s.) 1993 Blefuscuiana Iaculata Banner, Copestake
1986 Hedbergella aptica (Agalarova), Gorba- and White, p. 13, pI. 3, figs 1a-c, 2a-c.
chik, pp. 94, 234, pI. 14, figs 2, 3 (not figs 1993 Blefuscuiana praesimilis Banner, Cope-
4,5). stake and White, p. 14, pI. 8, figs 4a-c,
1988 Blefuscuiana aptica (Agalarova), Banner 5,6.
and Desai, p. 160, pI. 5, figs 4-7.
1993 Blefuscuiana infracretacea (Glaessner) Remarks: B. laculata has five subglobular
aptica (Agalarova), Banner, Copestake and chambers (hiL == 1 approx.) abutting the umbilicus,
White, p. 12, pI. 4, figs 4, 5. a wide umbilicus, a markedly lobulate periphery
1995 Hedbergella infracretacea (Glaessner), and intercameral sutural angles relatively narrow
Weiss, pI. 1, fig. 4. (about 120-135).
1996 Blefuscuiana infracretacea (Glaessner) Banner, Copestake and White (1993) described
occidentalis BouDagher-Fadel et al., p. identical forms as B. laculata and B. praesimilis
249, Fig. 1 (9-16). (respectively refigured in Plate 8.8, Figs 1-3 and
1997a Blefuscuiana infracretacea (Glaessner) Plate 8.9, Figs 1-3). As first revising authors we
occidentalis BouDagher-Fadel et al., pI. 1, choose the former as having priority.
figs 1-3.
Range: Early Barremian. Described from the
Remarks: B. infracretacea occidentalis has the Early Barremian of the Central North Sea.
same test shape, chamber shape and umbilical size
as B. infracretacea (Glaessner) s.s., but it has a Blefuscuiana laculata alobata Banner,
porticus and a smooth test surface and lacks per- Copestake and White, 1993
foration cones. Some forms have weak develop- (Plate 8.8, Figs 4-6; Figure 8.1)
ment of perforation cones on the first chamber of
the last whorl. They are transitional forms (Plate 1993 Blefuscuiana laculata alobata Banner,
8.7, Fig. 13) between the two subspecies. Copestake and White, p. 13, pI. 3, figs
3a-c, 4, 5a-c.
Range: (?Latest Barremian) Early Aptian- Late
Aptian. B. infracretacea occidentalis is slightly Remarks: This subspecies differs from B. Iacu-
older in stratigraphic range than B. infracretacea lata s.s. in having a less lobate periphery, a more
s .s. but it is not known to occur in the highest quadrate test and intercameral sutures relatively
Aptian or the Albian. It was not found in the broad (about 130-145).
Crimea, Azerbaijan, the Caucasus or North Africa,
but instead is known from the North Atlantic, NE Range: Early Barremian. First described from
England and the North Sea area (BouDagher-Fadel the Early Barremian of the Central North Sea.
et al., 1996 and 1997a). Weiss figured similar
Blefuscuiana mitra Banner and Desai, 1988
specimens from the Early and Middle Aptian of
Saxony. We have also identified specimens found (Plate 8.8, Figs 7, 8; Figure 8.1)
130 Blefuscuiana of the Praehedbergellidae

1959 ?Globigerina infracretacea Glaessner, 1990 Hedbergella occuZta Longoria; Weidich,

Obregon de la Parra, p. 149, pl. 3, figs p. 167, pl. 56, figs 16-18.
6a-b. 1995 Hedbergella cf. occulta Longoria; Weiss,
1988 Blefuscuiana mitra Banner and Desai, pl. I, figs 10-12.
p. 164, pl. 6, figs 4-6.
Remarks: This species has six sub globular
Remarks: This species has five chambers abut- chambers (hiL = 1 approx.) abutting the umbili-
ting the umbilicus. The chambers are normally cus, a flattened to slightly concave dorsal side,
subglobular, only the last ones are as high as long half of which occupied by early whorls, and,
(hiL = 1 approx.). The dorsal side is flat to ventrally, an open (uluh = 0.5 approx.) and deep
concave. The intercameral sutural angles are umbilicus.
narrow. Like B. speetonensis Banner and Desai
s.l., B. mitra shows a tendency towards plani- Range: Aptian. This species has previously
spirality in the last few chambers but it is distin- recorded from the Late Aptian of northern Mexico
guished by its smaller umbilicus (uluh = 0.4 and Speeton, North Yorkshire, England, and the
approx.) and fewer, more globular chambers (hiL Central North Sea; from the 'Middle' Aptian of
= 0.9 approx.). Bavaria and Germany, and from the Early Aptian
of the North Atlantic and the Central North Sea
Range: Late Aptian. This species was first area.
described from the Late Aptian Schackoina cabri
Blefuscuiana occulta (Longoria) perforocculta
Zone of the Speeton Cliff, North Yorkshire.
Banner, Copestake and White, 1993
Con specific forms were found by Obregon de la
Parra (1959) also from the Late Aptian cabri Zone (Plate 8.8, Figs 13, 14; Figure 8.1)
of Mexico.
1993 Blefuscuiana occulta (Longoria) perfor-
Blefuscuiana multicamerata Banner and Desai, occulta Banner, Copestake and White, pp.
1988 13, 14, pl. 8, figs 2a-c, 3.
(Plate 8.8, Figs 9, 10; Figure 8.1)
Remarks: This subspecies is characterized by
1988 Blefuscuiana multicamerata Banner and having perforation cones.
Desai, p. 165, pl. 6, figs 7a-d.
Range: Late Barremian to Late Aptian. It was
Remarks: This species has eight subglobular first described from the Central North Sea.
(hiL = 1 approx.) chambers in the last whorl. The
Blefuscuiana praetrocoidea (Kretchmar and
umbilicus (uluh = 0.5 approx.) is open and
Gorbachik, 1986)
shallow.
(Plate 8.9, Figs 4-12; Figures 8.1,8.2)
Range: Late Aptian. This species was first
1986 Hedbergella praetrocoidea Kretchmar and
described from the Upper Aptian Schackoina cabri
Gorbachik (in Gorbachik), p. 95, pl. 16,
Zone of Speeton Cliff, North Yorkshire.
figs 3-5.
Blefuscuiana occulta (Longoria) sensu stricto ?1993 Hedbergella praetrochoidea Gorbachik;
Shahin, pl. 6, fig. 4.
(Plate 8.8, Figs 11, 12; Figure 8.1)
1997a Blefuscuiana praetrocoidea (Kretchmar
1974 Hedbergella occuZta Longoria, pp. 63, 64, and Gorbachik), BouDagher-Fade1 et aZ.,
pl. 11, figs 7, 8; pl. 19, figs 17, 18; pl. 20, pl. I, figs 4-9, pl. 2, figs 1-10.
figs 5-9,17-18. 1997b Blefuscuiana praetrocoidea (Kretchmar
1988 Blefuscuiana occulta (Longoria), Banner and Gorbachik), BouDagher-Fadel et aZ.,
and Desai, p. 162, pl. 6, figs 8-12. pl. 3, figs 9, 10 (in press).
 Systematics 131

Remarks: B. praetrocoidea has six to seven came from the Kacha River of SW Crimea; it has
slightly depressed chambers visible dorsally in the not yet been recorded elsewhere.
last whorl, but only six abutting the umbilicus.
Blefuscuiana rudis Banner, Copestake and
The aperture extends fully to the periphery of the
White, 1993 sensu stricto
penultimate whorl, and possesses a broad portical
lip. The umbilicus is only slightly broad (uluh (Plate 8.10, Figs 4-6; Figure 8.1)
= 0.4 approx.).
This microperforate, non-muricate species 1993 Blefuscuiana rudis Banner, Copestake and
seems to be the ancestral form of the macroper- White (pars), p. 14, pI. 5, figs la--c (holo-
forate, muricate Hedbergella (BouDagher-Fadel et type only) (not paratype, figs 2, 5).
al., 1997a).
Through the courtesy of Dr T. Gorbachik we Remarks: A bulky test with marked dorsal
have been able to SEM paratypes of this species convexity and weakly depressed sutures. Early
from the Early Aptian of the Kacha River, SW chambers are subglobular (hiL = 1 approx.), but
Crimea (Plate 8, 9, Figs 4-12). they are predominantly depressed in the last whorl
(hiL = 0.7 approx.). There are five chambers in the
final whorl, all with strong perforation cones. The
Range: Early Aptian. This species has been
umbilicus is small and deep.
found with certainty only in the SW Crimea.
Shahin (1993, pI. 6, fig. 4) illustrated a specimen
under the name Hedbergella praetrochoidea (sic) Range: Barremian. The holotype was obtained
from the Late Aptian of Northern Sinai. His figure from the Central North Sea area (Early Barremian,
is not clear and the specimen is deformed, there- Lower Valhall Formation). It is not yet known
fore more information is needed to confirm its from elsewhere.
presence in this part of Tethys. Blefuscuiana rudis Banner, Copestake and
Blefuscuiana primare (Kretchmar and White pararudis subsp. nov.
Gorbachik, 1986) (Plate 8.10, Figs 7-9; Plate 8.11, Figs 1-3;
(Plate 8.10, Figs 1-3; Figure 8.1) Figure 8.1)

1986 Clavihedbergella primare Kretchmar and 1993 Blefuscuiana rudis Banner, Copestake and
Gorbachik (in Gorbachik), pp. 120, 121, pI. White (pars), p. 14, pI. 5, figs 2, 5 only.
25, figs 2, 3.
1997b Blefuscuiana primare (Kretchmar and Diagnosis: Morphologically the same as B. rudis
Gorbachik), BouDagher-Fadel et al., pI. 4, s.s. but there are six chambers in the last whorl.
figs 3,4 (in press).
Hoiotype: PF 52715, deposited in The Natural
Remarks: B. primare has a very small test of History Museum, London. From the Early Aptian
up to 170 j-Lm at its maximum diameter. Six of North Sea Well 20/2-2, 825 ft, Upper Valhall
slightly depressed chambers (hiL = 0.6 approx.) Formation.
abut the umbilicus, tending to increase gradually
in size. The dorsal side is flat to concave. The Range: Early Barremian to Early Aptian. This
umbilicus is shallow and very broad (uluh = 0.6 subspecies is known only from the Central North
approx.), with a narrow aperture and a narrow Sea area, where it ranges from the Lower to the
apertural rim. Upper part of the Valhall Formation.
Blefuscuiana speetonensis Banner and Desai,
Range: Late Barremian to Early Aptian. Both 1988 sensu stricto
the original material of Kretchmar and Gorbachik
(1986) and that of BouDagher-Fade1 et al. (1997b) (Plate 8.11, Figs 4-10; Figure 8.1)
132 Blefuscuiana of the Praehedbergellidae

1974 Hedbergella planispira (Tappan), Long- dorsal side with more curved intercameral sutures
oria, pp. 64, 65, pI. 23, figs 5-7,17,18. on the dorsal side and an aperture extending over
1988 Blefuscuiana speetonensis Banner and the periphery of the penultimate whorl with the
Desai, p. 164, pI. 7, figs 1-8. portici of the last three chambers encroaching onto
1993 Blefuscuiana speetonensis Banner and the dorsal side.
Desai; Shahin, p. 418, figs 5-10.
1997b Blefuscuiana speetonensis Banner and Range: Early Albian. B. speetonensis tunisiensis
Desai; BouDagher-Fadel et al., pI. 4, figs was first described from Tunisia. It derived in the
1, 2 (in press). Early Albian from the Late Aptian species spee-
tonensis s.s.
Remarks: B. speetonensis has six subglobular
Blefuscuiana sp.
chambers (hiL = 0.7 approx.) in the umbilicus,
and a flat to concave, evolute dorsal side. The (Plate 8.12, Figs 7-9; Figure 8.1)
umbilicus is moderately broad (uluh =0.5 approx.)
1986 Clavihedbergella tuschepsensis (Anton-
and open, with relict apertures and portici visible
ova), Gorbachik, pp. 118, 119,236, pI. 25,
for about half of the last whorl. The aperture has
fig. 1.
a proticus, which is broadest posteriorly, and
extends to the periphery of the penultimate whorl.
The umbilical shoulders of the chambers of the Remarks: When Antonova (1964) erected
last whorl are imperforate. Globigerina tuschepsensis, she published only
The tendency towards planispirality shown by drawings of her new species. Later, Gorbachik
this species suggests that it may be ancestral to (1986) used scanning electron micrographs to
multi-chambered species of Blowiella recorded illustrate a topotypic paratype with clearly asym-
from stratigraphically younger horizons of the metric chambers. However, she only illustrated a
Late Aptian. part of its dorsal side. We have now been able to
examine this paratype and photograph it by SEM
(Plate 8.12, Figs 7-9) to prove that it has indeed
Range: Youngest Early Aptian to early Late
more than four chambers abutting the umbilicus
Aptian. This species was first described from
and it belongs to the genus Blefuscuiana, while
North Yorkshire, England. Similar forms are
the holotype of tuschepsensis (also figured here by
recorded as H. planispira by Longoria from la
SEM for the first time on Plate 7.1, Figs 4--7) has
Drome, southeast France, and from Northern
only four chambers abutting the umbilicus, and is
Mexico.
a Praehedbergella. Blefuscuiana sp. is the only
Blefuscuiana speetonensis Banner and Desai known species of Blefuscuiana which possesses
tunisiensis BouDagher-Fadel, 1995 asymmetrical chambers, which become highest
posteriorly (hiL = 0.7 approx., but L'/L2 = 1.5
(Plate 8.12, Figs 1-3; Figure 8.1)
approx.).1t is known only from a single specimen
1995 Blefuscuiana speetonensis Banner and which was included amongst the two paratypes of
Desai tunisiensis BouDagher-Fadel, p. 144, Praehedbergella tuschepsensis by Antonova
pI. 2, figs 6, 7. (1964) in the collections of Moscow State
University.
Remarks: B. speetonensis tunisiensis also has
six chambers ventrally, in the last whorl. The last Range: This specimen was obtained from the
chambers on the dorsal side are almost as high as Early Barremian of the Tuscheps River (northwest
long (h/L =1 approx.). The aperture has a porticus Caucasus); it has not yet been recorded from
throughout its length, broadest posteriorly. anywhere else. As only one specimen is known,
B. speetonensis tunisiensis differs from B. spee- we prefer to leave for the present in open nomen-
tonensis speetonensis in having a more involute clature.
 Concluding comments 133

8.4 CONCLUDING COMMENTS last whorl (e.g. B. speetonensis Banner and Desai,
Plate 8.11, Figs 4-10, with six or seven chambers
Blefuscuiana directly evolved into Lilliputianella and B. multicamerata Banner and Desai, Plate 8.8,
in the latest Barremian (see Figure 5.2) by radial- Figs 9, 10, with up to eight, both Late Aptian
elongation of its later chambers (BouDagher- species with depressed dorsal spires and broad
Fadel, 1996), and L. globulifera (Kretchmar and umbilici). Another is the increase in the height of
Gorbachik) (Plate 9.1, Figs 5-15) is one species the spire to a maximum in the Early Aptian (e.g.
which shows that the evolution was gradual, with B. excelsa (Longoria) cumulus Banner et al.),
complete morphological transition. which was then followed by dorsally flat species
Blefusciana was also directly ancestral to which could even have concave dorsal sides (e.g.
Hedbergella (Figures 5.2, 8.2), which evolved B. speetonensis Banner and Desai, in the early
by gaining macroperforations and muricae (and by Late Aptian).
losing all possibility of developing perforation Just as in the case of species groups in Praehed-
cones). As we discuss in Chapter 13, it is probable bergella, the taxa of Blefuscuiana contain both
that the macroperforations allowed deeper waters species and subspecies, often distinguished by the
(at least, deeper than the surface of the waters) to presence or absence of perforation cones. In
become inhabited. Macroperforations were Blefuscuiana, when perforation cones were strongly
acquired first, and the muricae were only devel- developed, it was normal to lose the porticus or
oped later. For example, Hedbergella punctata aperturallip (e.g. B. infracretacea (Glaessner) s.s.,
Michael of the Late Albian possessed prominent B. aptiana (Bartenstein) orientalis BouDagher-
macroperforations but had not acquired muricae. Fadel et al., etc.). The related subspecies, which
Hedbergella pseudotrocoidea Michael, also of the lacked perforation cones, could develop the porticus
Late Albian, is a probable descendant of Blefus- or apertural lip (e.g. B. infracretacea (Glaessner)
cuiana gorbachikae (Longoria) (it is morphologi- occidentalis BouDagher-Fadel et al., B. aptiana
cally the same, with one additional chamber in the (Bartenstein) s.s.) (Figure 8.3). It is as though they
last whorl), and was conspicuously macroperforate could secrete only a limited amount of calcium
but still lacked muricae. It is probable that carbonate and that there was never enough for both
the evolution of Hedbergella occurred more than perforation cones and portici! However, a more
once. For example, Blefuscuiana praetrocoidea likely explanation could be that both the cones and
(Kretchmar and Gorbachik) was the immediate the portici were useful in food-grain disaggregation
ancestor to Hedbergella trocoidea (Gandolfi) and sorting prior to ingestion and that there was no
(Figure 8.2), the type species, and both macro- need for any subspecies to develop both. In contrast,
perforations and muricae appeared together in the subspecies of Praehedbergella (e.g. P. tuschepsen-
newly evolved species (BouDagher-Fadel et al., sis perforare Banner et al., P. ruka pap illata Banner
1996). The evolution of Hedbergella was almost et al.) could possess both perforation cones and
invariably accompanied by increase in test size. portici; this may have been because Praehed-
The Hedbergellidae were characteristically much bergella species were smaller than the smallest
larger than the ancestral Praehedbergellidae. Blefuscuiana, so that their areas for food-grain dis-
Two more trends of evolution are apparent. One aggregation were also smaller and both tools were
is the increase in the number of chambers in the needed.
132 127 121 112 Ma

APTIAN ALBIAN
HAUTERIVIAN BARREMIAN
EARLY LATE EARLY LATE

B. infracretacea occidentalis
\
\

B. praetrocoidea
\
\
\
H. trocoidea
,
\
\
\
\ T. roberti

Figure 8.2 The stratigraphical relationship between Blefuscuiana infracretacea (Glaessner) occidentalis BouDagher-Fadel et al., 1996, B. praetrocoidea (Kretchmar and
Gorbachik, 1986), Hedbergella trocoidea (Gandolfi, 1942) and Ticinella roberti (Gandolfi, 1942) (from BouDagher-Fadel et at., I 997a).
 127 121 112.2 Ma

BARREMIAN APTIAN ALBIAN

EARLY LATE EARLY LATE EARLY LATE

B. infracretacea 5.5.
,,
: B. infracretacea occidentalis
\
\
I.
\ , B. h/se. amae

. I
B. apt/ana s.s.
\
\ \
\ \
\ \
\ B. albiana
\
\
\
\
\ B. aptiana orienta/is

Figure 8.3 The evolution of Blefuscuiana infracretacea (Glaessner) sensu lato and B. aptiana (Bartenstein) sensu lato in the Early Cretaceous (adapted from BouDagher-
Fadel et al., 1996, Fig. 5).
Plate 8.1

Figures 1-5 Blejuscuiana kuznetsovae Banner and Desai, 1988. Late Aptian, Schackoina cabri Zone, Speeton Cliff, NE England,
deposited in The Natural History Museum, London, nos PF 52091 (holotype), PF 52093, PF 52094, PF 52096
(paratypes): 1, 2, ventral side of holotype (x175) and detail of surface of last chamber showing a microperforate
and smooth wall (x1800); 3-5, ventral, dorsal and peripheral views of three paratypes (x165, x195 and x195,
respectively).
Figures 6-9 Blejuscuiana albiana BouDagher-Fadel et al., 1996. Late Albian, sample N149, Predoroshnaya, SW Crimea, holo-
type deposited in The Natural History Museum, London, no. PF 53012: 6,9, ventral view of paratype (x200) and
enlargement of the surface showing microperforate wall with a tendency to larger pores (but still irregularly spaced)
(x1200); 7,8, dorsal and ventral views of hoiotype (x175).
Plate 8.2

Figures 1-8 Blejuscuiana aptiana (Bartenstein, 1965) sensu stricto. 1-4, Early Aptian, Mittelland-Kanal, near Wenden,
Braunschweig, central Germany, deposited in the Senckenberg Museum, Frankfurt-on-Main, Germany, nos SMF
(27) 2346a-<l (from Bartenstein, 1965, text-figs 3-6): 1, ventral view of holotype (xI00); 2-4, dorsal, ventral and
dorsal views of paratypes (xl00). 5-8, Late Aptian, upper Schackoina cabri Zone, Speeton Cliff, NE England,
deposited in The Natural History Museum, London, nos PF 52082-PF 82084: 5-7, dorsal (x240), peripheral
(x240) and ventral views (xI80); 8, enlargement of wall surface showing microperforations (xI500).
Figures 9-11 Blefuscuiana aptiana (Bartenstein) orientalis BouDagher-Fadel et al., 1996. Form with perforation cones in the
early whorls, high chambers in the last whorl and a slightly concave dorsal side. Early Aptian, sample AGA-l,
near Nardaran village, NE Azerbaijan, deposited in The Natural History Museum, London, nos PF 53013 (holo-
type), PF 53020 (paratype): 9, 10, ventral and dorsal views of holotype (xI65); 11, ventral view of paratype
(xl90).
Plate 8.3
Figures 1-8 Blefuscuiana aptiana (Bartenstein) orientalis BouDagher-Fadel et al., 1996, 1,2, Early Aptian, sample AGA-l,
near Nardaran village, NE Azerbaijan, paratype deposited in The Natural History Museum, London, no. PF 53017:
ventral view (x205), and enlargement of the surface showing clearly the perforation cones (xI400). 3-8, Aptian,
Krasnaya, SW Crimea, paratypes deposited in The Natural History Museum, London, nos PF 53015, PF 53016:
3,5, ventral and dorsal views (xI65), 4, enlargement of the wall surface showing very weak perforation cones in
the penultimate chamber of umbilical side (perforation cones are present on all earlier chambers) (x860); 6, 8,
dorsal and ventral views (x165), 7, enlargement of the wall surface showing very weak perforation cones in the
penultimate chamber of umbilical side (perforation cones are present on all earlier chambers) (x865).
Plate 8A

Figures 1-8 Blefuscuiana convexa (Longoria, 1974). 1-3, Early Late Aptian, Schackoina cabri Zone, la Dr6me region, SE
France (from Longoria, 1974, pI. 14, figs 12-14), holotype deposited in the US National Museum of Natural
History, Washington, DC, no. 185019: dorsal, peripheral and ventral views (x330). 4-6, (= Loeblichella moulladei
Longoria, 1974), early Late Aptian, Schackoina cabri Zone, la Drome region, SE France (from Longoria, 1974,
pI. 14, figs 15-17), holotype deposited in the US National Museum of Natural History, Washington, DC, no.
185020: dorsal, peripheral and ventral views (x290). 7, 8, earliest Late Aptian, Hluck Formation, Outer Flysch
Belt (Magura group of nappes), West Carpathians, deposited in collection of the Czech Geological Survey, Bmo:
dorsal and peripheral views (x280).
Plate 8.5

Figures 1-3 Blefuscuiana daminiae Banner, Copestake and White, 1993. Early Aptian, Well 15/30-3, 12830-40 ft, central
North Sea, holotype deposited in The Natural History Museum, London, no. PF 52696: peripheral, ventral and
dorsal views (x215).
Figures 4-6 Blefuscuiana excelsa (Longoria, 1974) sensu stricto. Early Aptian, la Drome region, SE France (from Longoria,
1974, pI. 18, figs 14-16), holotype deposited in the US National Museum of Natural History, Washington DC,
no. 184998: ventral, peripheral and dorsal views (x280).
Figures 7-12 Blefuscuiana excelsa (Longoria) cumulus Banner, Copestake and White, 1993.7-9, Early Aptian, Well 15/30-3,
12830-40 ft, central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52708:
ventral, peripheral and dorsal views (x250); 10-12, Early Aptian, sample N26, Kacha River, Crimea, deposited
in The Natural History Museum, London, nos PF 53074, PF 53076, PF 53077: 10, dorsal view of a very high
spired form (xI90); 11, ventral view showing a small umbilicus and a narrow slit aperture bordered with a porticus
(x335); 12, dorsal view (x230).
Plate 8.6

Figures 1-6 Blefuscuiana gorbachikae (Longoria, 1974). 1-3, Late Aptian, La Boca Canyon, Sierra de la Silla, SSE of
Monterrey, Mexico (from Longoria, 1974, pI. 15, figs 11-13), holotype deposited in US National Museum
of Natural History, Washington, DC, no. 185000: ventral, peripheral and dorsal views (x 170). 4, Late Aptian,
sample N120, Kacha River, SW Crimea, deposited in The Natural History Museum, London, no. PF 53078:
ventral view (x175). 5, 6, Early Aptian, SS354, Jebel Chenanrafa, Tunisia: ventral view (with a strongly over-
hanging last chamber) and dorsal view (x230).
Figures 7, 8 BI~fuscuiana hexacamerata sp. nov. Late Aptian, sample N120, Kacha River, SW Crimea, holotype deposited in
The Natural History Museum. London, no. PF 53079: dorsal and ventral views (x200).
Figures 9-11 Blefuscuiana hispaniae (Longoria, 1974). Late Aptian, Schackoina cabri Zone, La Boca Canyon, Sierra de la
Silla, SSE of Monterrey, Mexico (from Longoria, 1974, pI. 19, figs 6-8), holotype deposited in the US National
Museum of Natural History, Washington, DC, no. 185002: dorsal, peripheral and ventral views (x21O).
Plate 8.7

Figures 1-8 Blefuscuiana infracretacrea (Glaessner, 1937) sensu stricto. 1-5, Albian, Ilkaya, North Caucasus, metatypes
collected by Glaessner in 1935 (SEM photography by the Environmental Chamber technique at The Natural
History Museum, London), deposited in the US National Museum of Natural History, Washington, DC, slide no.
689757: 1, 3, dorsal and ventral views (xI80), 2, perforation cones on ventral side of second chamber of last
whorl (x600): 4, 5, ventral and dorsal views (xI95). 6--8, latest Aptian sample OG60, Otto Gott pit, Sarstedt,
south of Hannover, Germany (figured by Banner and Desai, 1988, pI. 3, figs 4, 5 as B. cf. aptica (Agalarova,
deposited in The Natural History Museum, London, nos PF 52102, PF 52103: 6, ventral view (x250), 7, dorsal
view showing close perforation-cones in the early whorls (x265), 8, penultimate chamber showing detailed
construction of the perforation cones (x1140).
Figures 9-15 Blefuscuiana infracretacea (Glaessner) occidentalis BouDagher-Fadel et al., 1996.9-11, Early Aptian, North Sea
Well 2012-2, 8300 ft, holotype deposited in The Natural History Museum, London, no. PF 52102: ventral, dorsal
and peripheral views (xI95). 12-15, Late Aptian, sample 8130, Schackoina cabri Zone, Speeton Cliff, NE England,
paratypes deposited in The Natural History Museum, London, nos PF 52098-PF52101: 12, enlargement of the
surface showing a finely perforated wall (xI5OO); 13, 15, two ventral views showing clearly the thin porticus
(x225); 14, dorsal view (x245).
Plate 8.8

Figures 1-3 Blefuscuiana laculata Banner, Copestake and White, 1993 sensu stricto. Early Barremian, Well 20/2-2, 8680
m, central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52724: dorsal, periph-
eral and ventral views (x255).
Figures 4-6 Blefuscuiana laculata alobata Banner, Copestake and White, 1993. Early Barremian, Well 20/2-2, 8680 m,
central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52726: dorsal, periph-
eral and ventral views (x240).
Figures 7, 8 Blefuscuiana mitra Banner and Desai, 1988. Earliest Late Aptian, Schackoina cabri Zone, Speeton Cliff, NE
England, deposited in The Natural History Museum, London, nos PF 52089 (holotype), PF 52088 (paratype):
7, dorsal view of holotype (xI60); 8, peripheral view of paratype (x200).
Figures 9, 10 Blefuscuiana multicamerata Banner and Desai, 1988. Late Aptian, Schackoina cabri Zone, sample 8103, Speeton
Cliff, NE England, holotype deposited in The Natural History Museum, London, NHML PF52090: Dorsal and
ventral views (x400).
Figures 11, 12 Blefuscuiana occulta (Longoria, 1974) sensu stricto. Late Aptian, Schackoina cabri Zone, La Pena Formation,
La Boca Canyon, Mexico (from Longoria, 1974, pI. 20, figs 8, 9), holotype deposited in the US National
Museum of Natural History, Washington, DC, no. 185011: dorsal and ventral views (xI60).
Figures 13, 14 Blefuscuiana occulta (Longoria) perforocculta Banner, Copestake and White, 1993. Early Aptian, Well 20/2-2,
8300 ft, central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52678: periph-
eral and ventral views (xI80).
Plate 8.9

Figures 1-3 Blefuscuiana laculata Banner. Copestake and White. 1993 sensu stricto (= Blefuscuiana praesimilis Banner,
Copestake and White, 1993). Early Aptian. Well 20/2-2, 8300 ft, central North Sea, holotype deposited in The
Natural History Museum, London, no. PF 52691: Dorsal, peripheral and ventral views (x230).
Figures 4-12 Blefuscuiana praetrocoidea (Kretchmar and Gorbachik, 1986). Early Aptian, Kacha River, SW Crimea, four
paratypes deposited in the Geological Faculty, Moscow State University, Russia, photographed at The Natural
History Museum, London: 4, 7, 9, 12 SEM using the Environmental Chamber technique (without coating): 4,
dorsal view (x240); 7, 9, 12, ventral, peripheral and dorsal views (x235); 5, 6, peripheral and ventral views
(x225); 8, 10, 11, oblique-peripheral, ventral and peripheral views (x260).
Plate 8.10

Figures 1-3 Blefuscuiana primare (Kretchmar and Gorbachik, 1986), I, 2, Late Aptian, sample N120, Kacha River, SW Crimea,
topotype deposited in The Natural History Museum, London, no. PF 62912: dorsal and ventral views (x300).
3, Late Aptian, Kacha River, SW Crimea, topotype (from Gorbachik, 1986, pI. 25, fig. 2a), deposited in the
Department of Palaeontology, Moscow State University, Russia: dorsal view (x460).
Figures 4--6 Blefuscuiana rudis Banner, Copestake and White, 1993 sensu stricto. Early Barremian, Well 16/28-6RE, 4275 m,
central North Sea, holotype deposited in The Natural History Museum, London, no. PF 52713: ventral, peripheral
and dorsal views (xI80).
Figures 7-9 Blefuscuiana rudis Banner, Copestake and White pararudis subsp. nov. Early Aptian, central North Sea, Well
20/2-2, 8250 ft, holotype deposited in The Natural History Museum, London, no. PF 52715: ventral, peripheral
and dorsal views (x375).
Plate 8.11

Figures 1-3 Blefuscuiana rudis Banner. Copestake and White pararudis sUbsp.nov. Early Barremian, Well I 6/28-6RE ,
4195 m, central North Sea, paratype deposited in The Natural History Museum, London, no. PF 52714: dorsal,
peripheral and ventral views (x31O).
Figures 4-10 Blefuscuiana speetonensis Banner and Desai, 1988 sensu stricto. 4-7, early Late Aptian, Schackoina cabri Zone,
sample 8109, Speeton Cliff, NE England, deposited in The Natural History Museum, London, nos PF 52075
(holotype), PF 52078, PF 52080, PF 52081 (paratypes): 4, dorsal view of holotype (x200); 5,6,7, ventral (xI85),
dorsal (x21O) and peripheral (x230) views of three paratypes. 8-10, Late Aptian, sample N120, Kacha River, SW
Crimea, deposited in The Natural History Museum, London, nos PF 53084, PF 53085: 8, 9, ventral and dorsal
views (xI90); 10, dorsal view (xI50).
Plate 8.12

Figures 1-3 Blefuscuiana speetonensis Banner and Desai tunisiensis BouDagher-Fadel, 1995. Early Albian, SS538, Beauvoir-
VI Well, Tunisia, deposited in The Natural History Museum, London: 1, holotype, dorsal side almost completely
involute and concave (xI55); 2, 3, paratypes, dorsal view (xI55) and ventral view with a large umbilicus and an
aperture going right over the periphery (xI73).
Figures 4-6 Blefuscuiana daminiae Banner, Copestake and White, 1993. (= Blefuscuiana whittakeri Banner, Copestake and
White, 1993), Early Aptian, Well 15/30-3, 12830-12840 ft, central North Sea, deposited in The Natural History
Museum, London, no. PF 52717: dorsal, peripheral and ventral views (x290).
Figures 7-9 Blefuscuiana sp. Early Barremian, Tuscheps River, northwest Caucasus, deposited in the Departruent of Geology,
Moscow State University, Russia: ventral, peripheral and dorsal views (x425).
 9
The Praehedbergellidae
with elongate chambers
- Lilliputianella,
Lilliputianelloides gen.nov.,
and Wondersella
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

9.1 INTRODUCTION different stratigraphic ranges (Figure 9.1) and

different geographic distributions. Lilliputian-
As we discuss further in Chapter 13, the ability elloides is confined to the Barremian and is
of the foraminifera to thrive under oxygen concen- known, so far, only from north Africa and south-
trations less than saturation would be enhanced by eastern Europe. Lilliputianelia occurred only
enabling the organism to increase the surface area from the latest Barremian through to the end of
of its chambers. The simple way to do this would the Aptian and is known from Central America,
be to elongate the chambers in a radial direction. northern and southern Europe and north Africa;
The globular or slightly depressed chambers of it may yet prove to have had a worldwide distri-
Blefuscuiana did, in fact, become elongate. These bution.
elongated species, still trochospiral in coiling Chamber elongation also occurred in the
mode, are united by us into the genera Lilli- latest Aptian of the Middle East in Wondersella,
putianelia and Lilliputianelioides gen.nov. The a genus which was also characterized by strepto-
genus Lilliputianella clearly evolved from Blefus- spiral coiling in the final stages of growth.
cuiana in the latest Barremian, while Lilliputian- This morphology is also discussed further in
elloides derived directly from Praehedbergella in Chapter 13.
the later part of the Early Barremian (Figure 5.2).
These two genera have different morphologies,
 .... .... ....
....
N N N
..... .... ~ f
BARREMIAN APTIAN ALBIAN
C)
m
EARLY LATE EARLY LATE EARLY LATE

Lllilputianella blzonae
L. globullfera

-- L. kuhryl
L. /abocaensls
L./ongorl/

L. rob/esse
Lllliputianelloides eocretaceus
1--- Wondersella athersuchl
-

Figure 9.1 Stratigraphical distribution of Early Cretaceous Lilliputianella, Lilliputianelloides gen.nov. and Wondersella. Chronostratigraphy and Absolute Time Scale after
Gradstein et al. (1995).
 Systematics 163

9.2 SYSTEMATICS putianella longorii Banner and Desai,

1988).
Superfamily GLOBIGERINACEA Carpenter, non 1994 Clavihedbergella eocretacea (Neagu) ,
Parker and Jones, 1862 Coccioni and Premoli Silva, figs 9,
10-18 (rect. L. Zongorii pars.).
Family PRAEHEDBERGELLIDAE Banner
and Desai, 1988 Remarks: L. eocretaceus has four chambers
abutting the umbilicus in the last whorl. Its last
Genus Lilliputianelloides gen.nov.
chambers are broadly elongate. The umbilicus is
Type species CZavihedbergella eocretacea very narrow and the aperture extends from the
Neagu, 1975 umbilicus to the periphery of the penultimate
whorl and is furnished with a narrow lip. The
Diagnosis: A praehedbergellid; trochospiral, sutural intercameral angles are very broad before
with an evolute dorsal side and an involute ventral the chambers become elongate; with the elonga-
side. Wall microperforate and smooth. Aperture tion these angles become more narrow.
interio-marginal extending from the umbilicus The type species (holotype and all of its
toward the periphery of the penultimate whorl; paratypes) distinctly shows an initial test of
intra-extraumbilical. Four chambers only visible in PraehedbergeZla form. The test is smooth and
the last whorl on the umbilical side and only the microperforate (Plate 9.1, Figs 3, 4) so it is not
last four chambers abutting the umbilicus. Later a CZavihedbergella as Neagu (1975) supposed.
chambers radially elongate and higher than broad. Only the last one or two chambers show radial
[Gender: masculine.] elongation.
So far, the genus is monotypic, therefore the
Remarks: Lilliputianelloides has only four characters of the genus are those of the type
chambers both in the final whorl and abutting the species.
umbilicus and only the last one or two chambers
tend to show radial elongation. Lilliputianella has Range: Latest Early Barremian-Late Barremian.
five or more chambers abutting the umbilicus and, The type specimens came from the Carhaga beds
typically, the chambers are much more radially (Late Barremian) of the Izvorul Valley, Romania.
elongate. Both genera have a different evolu- The species has also been found in the latest Early
tionary history (Lilliputianelloides has an initial to Late Barremian of Tunisia. It has not yet been
test of Praehedbergella type), and different strati- found in Aptian or younger strata.
graphic and geographic ranges (Introduction, Genus Lilliputianella Banner and Desai, 1988
above).
Type species Lilliputianella Zongorii Banner and
Desai, 1988
Range: Latest Early Barremian-Late Barremian.
Lilliputianelloides eocretaceus (Neagu, 1975)
Key to the species of Lilliputianella
(Plate 9.1, Figs 1-4; Figure 9.1)
1 Five to six chambers in the last whorl,
1975 CZavihedbergella eocretacea Neagu, chambers are higher than long but are not
pp. 112, 113; pI. 89, figs 1-6, 9, 10 elongate
(holotype), text-fig. 20. 1.1 Umbilicus broad L. globulifera
1995 Lilliputianella sp. 1, BouDagher- (= Hedbergella maslakovae
Fadel, p. 140. =
Hedbergella similis)
non 1993 Lilliputianella eocretacea (Neagu), 1.2 Umbilicus narrow L. labocaensis
Banner, Copestake and White, p. 15, 2 Five to six chambers in the last whorl, very
pI. 6, fig. 8, pI. 8, fig. 7 (rect. Lilli- high and radially elongated
164 The Praehedbergellidae with elongate chambers

2.1 Early chambers of the last whorl peripher- 1974 Hedbergella bollii Longoria, p. 53, pI. 13,
ally rounded, only last two or three are figs 1, 2, 6-8, 10-14.
high but not very elongate 1986 Clavihedbergella bollii (Longoria),
2.1.1 Umbilicus narrow L. longorii Gorbachik, pp. 106, 107, pI. 26, figs 4-6.
2.1.2 Umbilicus broad L. kuhryi 1988 Lilliputianella bizonae (Chevalier), Banner
2.2 Early chambers of last whorl peripherally and Desai, p. 168, pI. 7, figs 12-15.
rounded but the last two or three are radi- 1993 Lilliputianella bizonae (Chevalier), Shahin,
ally very elongate L. roblesae p.422.
2.3 All chambers of the last whorl are higher 1995 ?Globigerinelloides bizonae (Chevalier),
than long and most or all are strongly radi- Weiss, pI. 2, figs 3-5.
ally elongate L. bizonae 1997b Lilliputianella bizonae (Chevalier),
(= Hedbergella bollii) BouDagher-Fadel et aZ., pI. 5, figs 5,6, pI.
6, figs 6-9.
Lilliputianella longorii Banner and Desai, 1988
(Plate 9.2, Figs 4-6; Figure 9.1) Remarks: This species is characterized by
having 5)1 to 6 chambers in the last whorl, which
1988 Lilliputianella Zongorii Banner and Desai,
become radially elongated and broadly pointed,
pp. 165-168, pI. 4, figs 11, 12.
and a broad, shallow umbilicus in which relict
1993 Lilliputianella eocretacea (Neagu), Banner,
apertures of at least three chambers are visible.
Copestake and White, p. 15, pI. 6, fig. 8,
The type specimens of bizonae and bollii are
pI. 8, figs 7a-c (non Clavihedbergella
identical (our Plate 9.3, Figs 8-10 and Figs 15-17,
eocretacea Neagu, 1975). respectively), and are therefore considered here to
1994 CZavihedbergelia eocretacea Neagu;
be synonymous.
Coccioni and Premoli Silva, pp. 669,670,
figs 9,10-18 (non Neagu, 1975). Range: Late Aptian. Chevalier (1961) described
his species from the early Late Aptian of France.
Remarks: This species has five to six chambers The specimens described by Gorbachik (1971,
in the last whorl; the early periphery is rounded 1986) and BouDagher-Fadel et aZ. (1997b) came
but the last two or three chambers increase rapidly from the Late Aptian of the Crimea. Longoria
in height to become one and a half to twice as (1974) illustrated his type material from the Late
high as long in umbilical view, and ovoid in form, Aptian of la Drome, France and from La Pena
being bluntly rounded or conical at their distal Formation and La Boca Canyon in Mexico.
ends. The umbilicus is deep but narrow, in breadth Banner and Desai (1988) recorded similar forms
being about 10-12% of the maximum test diam- from the Schackoina cabri Zone of Speeton Cliff
eter. The aperture is in an umbilical-extraumbilical in England as did Weiss (1995) from the same
slit, furnished with a narrow porticus. zone of the Wiechendorf 1/86 Borehole of Lower
Saxony. Shahin (1993) recorded L. bizonae from
Range: Late Aptian. L. Zongorii was first the early part of the Late Aptian of Egypt.
described from the Late Aptian, Schackoina cabri
Zone of Speeton, England. Other records are also Lilliputianella globulifera (Kretchmar and
Late Aptian. Gorbachik, 1971)

Lilliputianella bizonae (Chevalier, 1961) (Plate 9.1, Figs 5-15; Figure 9.1)
1971 Clavihedbergella globulifera Gorbachik
(Plate 9.3, Figs 8-17; Figure 9.1)
and Kretchmar, p. 22, pI. 6, figs 6, 7
1961 Hastigerinella bizonae Chevalier, p. 34, pI. (nomen nudum).
1, figs 24-28. 1971 Clavihedbergella globulifera Kretchmar
1971 Clavihedbergella bizonae (Chevalier), and Gorbachik (in Gorbachik), p. 136, pI.
Gorbachik, p. 138, pI. 30, figs 4a-b. 30, figs 1a-c (holotype).
 Systematics 165

1974 Hedbergella maslakovae Longoria, pp. three become radially ovoid, pointed and laterally
61-63, pI. 20, figs 1-3, 14--16, pI. 24, figs compressed. The umbilicus is broad and shallow
11-14. with a diameter equal to as much as a third of that
1974 Hedbergella similis Longoria, pp. 68, 69, of the whole test.
pI. 16, figs 10-21, pI. 18, figs 12, 13,
pI. 23, figs 14--16. Range: Late Aptian (?later part of the Early
1988 Lilliputianella globulifera (Kretchmar and Aptian). Longoria (1974) believed L. kuhryi to be
Gorbachik), Banner and Desai, p. 168, restricted to the upper part of the Schackoina cabri
pI. 8, figs 1-4. Zone. Salaj's specimens, which are doubtfully
1997b Lilliputianella globulifera (Kretchmar and referable to this species, were obtained from
Gorbachik), BouDagher-Fadel et al., pI. 6, Tunisian strata referred to the basal Bedoulian
fig. 5. (later part of the Early Aptian). Banner and
Desai's specimens are also Late Aptian, belonging
Remarks: This species is distinguished by having to the upper part of the S. cabri Zone.
5~ to 7 chambers in the last whorl, the last two
Lilliputianella labocaensis (Longoria, 1974)
chambers are distinctly higher than long, but are not
elongate. The umbilicus is largely broad and deep. (Plate 9.2, Figs 1-3; Figure 9.1)
The original illustrations of the synonyms 1974 Hedbergella labocaensis Longoria, pp. 60,
Hedbergella maslakovae and H. similis (both 61, pI. 16, figs 7-9, 22-24.
described by Longoria, 1974) are shown in Plate
9.1, Figs 10-14. Remarks: This species has five to six chambers
in the last whorl, the last two or three being higher
Range: Latest Barremian-Aptian. Kretchmar than long but not elongate. Umbilicus shallow and
and Gorbachik obtained the primary types from narrow.
the Aptian of the Crimea and they were recorded
by Gorbachik (1986) and by BouDagher-Fadel et Range: Late Aptian. Longoria described his
ai. (1997b) from the Aptian of the same area. species from the La Pefia Formation, La Boca
Banner and Desai (1988) reported this species Canyon, Mexico, and from SE France.
from the early Late Aptian (Schackoina cabri
Zone) of Speeton Cliff, NE England, Shahin Lilliputianella roblesae (Obregon de la Parra,
(1933) from the latest Barremian-Early Aptian of 1959)
Egypt. Longoria's (1974) records of Hedbergella (Plate 9.3 Figs 1-7; Figure 9.1)
similis and H. masiakovae came from the Late
Aptian of the La Pefia Formation, La Boca 1959 Giobigerina roblesae Obregon de la Parra,
Canyon, Mexico. p. 149, pI. 4, figs 10, 11.
1974 Hedbergella robiesae (Obregon), Longoria,
Lilliputianella kuhryi (Longoria, 1974) pp. 65, 66, pI. 16, figs 1-3,4--6, pI. 20, figs
(Plate 9.2 Figs 7-10; Figure 9.1) 10,11.
1997b Lilliputianella roblesae (Obregon),
1974 Hedbergella kuhryi Longoria, p. 60, pI. 14, BouDagher-Fadel et al., pI. 5, figs 5,6.
figs 1--6.
?1984 Clavihedbergella kuhryi (Longoria), Salaj, Remarks: This species has strongly elongate
p. 594, pI. 3, fig. 13. last three chambers and a broad umbilicus.
1988 Lilliputianella kuhryi (Longoria), Banner
and Desai, p. 169, pI. 8, fig. 5. Range: Late Aptian. L. roblesae was originally
described from the La Pefia Formation of northern
Remarks: This species has six chambers in the Mexico. It has also been reported by Chevalier
final whorl; chambers are subspherical, but the last (1961) from the Early Gargasian of La Bectoule,
166 The Praehedbergellidae with elongate chambers

France, and from the Schackoina cabri Zone of suchi extends upwards into the lowest Nahr Umr
Spain, France and Mexico (Longoria, 1974) and Formation, Albian, but we think this is improb-
from the Late Aptian of the Crimea (BouDagher- able.
Fadel et al., 1997b).
Genus Wondersella Banner and Strank, 1987
9.3 CONCLUDING COMMENTS
Type species Wondersella athersuchi Banner
and Strank, 1987
The elongation of the chambers in lilliputian-
Wondersella athersuchi Banner and Strank, 1987 elloides (derived from Praehedbergella in the
Early Barremian; Figure 5.2), Lilliputianella and
(Plate 9.4 Figs 1-7; Figure 9.1)
Wondersella (both independently from species of
1987 Wondersella athersuchi Banner and Blefuscuiana, the former in the latest Barremian
Strank, pp. 39-48, pI. 1, figs 1-9, pI. 2, and the latter in the latest Aptian) was probably
figs 1-10, pI. 3, figs 1-14, pI. 4, figs 1-6; associated with reduced oxycity, as suggested and
text-figs 3a-c. discussed in Chapter 13. The same may have
1988 Wondersella athersuchi Banner and happened with Asterohedbergella in its Israeli
Strank, p. 186. Late Cenomanian occurrence, which might have
been contemporaneous with the second anoxic
Remarks: A praehedbergellid in which the period of the proto-Atlantic and western Tethys.
early trochospiral test is followed by a strep- Nevertheless, none of these taxa had any descen-
tospire. The first two whorls are very similar to dants. None of the Praehedbergellinae with
those of Blefuscuiana gorbachikae (Longoria, broadly elongate chambers survived the Aptian
1974; Chapter 8) but the succeeding chambers are (except possibly Wondersella, which may have
crescentiform (lunate) in equatorial view and survived into the earliest Albian).
are coiled irregularly and streptospirally, Lilliputianella has been, in the past, confused
frequently covering and closing the ventral with Clavihedbergella, because both are Creta-
umbilicus. ceous and have trochospiral coils with radial
It has similarity to Asterohedbergella astero- chamber elongation. However, the wall surfaces
spinosa Hamaoui, 1964, from the Late Ceno- were very different, Lilliputianella being smooth
manian of Israel, but that species is a hedbergellid and microperforate like its ancestral Blefuscuiana,
and does not have the lunate chambers or strepto- while Clavihedbergella (which first occurred in
spirality of Wondersella. the Late Albian and became common in the
Described originally from thin-section material. Cenomanian) was macroperforate, muricate and
The genus is monotypic so the characters of the larger, and arose from Hedbergella. The genus
genus are those of the type species. This taxon has Clavihedbergella was originally proposed by
no descendants. Banner and Blow (1959), who noticed that it first
evolved in the Late Albian; Loeblich and Tappan
Range: Latest Aptian (?earliest Albian). This (1964) attempted to correct this, by claiming that
species was first described from the highest they had many specimens from the Aptian. Of
Shuaiba Formation (Late Aptian) of offshore wells course, they had seen many, many Aptian speci-
in the United Arab Emirates, Abu Dhabi. At these mens of Lilliputianella!
localities it is possible that the range of W. ather-
Plate 9.1

Figures 1-4 Lilliputianelloides eocretaceus (Neagu, 1975). 1,2, Late Barremian, Carhaga beds, Izvorul valley, Romania (from
Neagu, 1975, pI. 89, figs 9, 10), holotype deposited in the collections of the Laboratory of Palaeontology, University
of Bucharest, no. 10.288: dorsal and ventral views (xl60). 3,4, Late Barremian, SS94, Beauvoir-I Well, Tunisia
deposited in The Natural History Museum, London, PF 62923: ventral and dorsal views (xI85).
Figures 5-15 Lilliputianella globulifera (Kretchmar and Gorbachik, 1971). Late Aptian, the Crimea (from Kretchmar and
Gorbachik, 1971, pI. 30, figs la-<:) , holotype deposited in the Geology Department, Moscow State University,
no. 121-72: 5-7, dorsal, peripheral and ventral views (xI25); 8,9, Late Aptian, Krasnaya, SW Crimea, topotype
deposited in The Natural History Museum, London, no. PF 62913: dorsal (xI85) and ventral (x245) views. 10,
11, (= Hedbergella maslakovae Longoria, 1974), Late Aptian, La Boca Canyon, Sierra de la Silla, Mexico (from
Longoria, 1974, pI. 24, figs 11, 12), holotype deposited in the US National Museum of Natural History, Washington,
DC, no. 185009: dorsal and ventral views (xI55). 12-14, (= Hedbergella similis Longoria, 1974), earliest Late
Aptian, Schackoina cabri Zone, La Pena Formation, La Boca Canyon, Sierra de la Silla, Mexico (from Longoria,
1974, pI. 16, figs 16-18), holotype deposited in the US National Museum of Natural History, Washington, DC,
no. 185017: dorsal, peripheral and ventral views (xl60); 15, early Late Aptian, sample 8109, Schackoina cabri
Zone, Speeton Cliff, NE England (figured by Banner and Desai, 1988, pI. 8, fig. 9a, as Lilliputianella similis
(Longoria)), deposited in The Natural History Museum, London, PF 52118: ventral view (x21O).
Plate 9.2

Figures 1-3 Lilliputianella labocaensis (Longoria, 1974). Late Aptian, La Boca Canyon, Sierra de la Silla, Mexico (from
Longoria, 1974, pI. 16, figs 22-24), holotype deposited in the US National Museum of Natural History, Washington,
DC, no. 185006: dorsal, peripheral and ventral views (xI65).
Figures 4-6 Lilliputianella longorii Banner and Desai, 1988. Earliest part of the Late Aptian, Schackoina cabri Zone, sample
8103, Speeton Cliff, NE England, deposited in The Natural History Museum, London, nos PF 52116 (holotype),
PF 52117 (paratype): 4, 5, ventral and dorsal views of holotype (x200); 6, ventral view of paratype (x21O).
Figures 7-10 Lilliputianella kuhryi (Longoria, 1974).7-9, Late Aptian, upper Schackoina cabri Zone, Argos Formation, Murica
Province, SE Spain (from Longoria, 1974, pI. 14, figs 1-3), paratype: dorsal, ventral and peripheral views (xl60).
10, earliest Late Aptian, upper Schackoina cabri Zone, Speeton Cliff, NE England, deposited in The Natural
History Museum, London, no. PF 52124: ventral view (x270)
Plate 9.3

Figures 1-7 Lilliputianella roblesae (Obreg6n de 1a Parra, 1959). Late Aptian, La Pena Formation, Mexico (from Obreg6n de
la Parra, 1959, pI. 4, figs 4a, b), holotype (depository not given): ventral and dorsal views (x95). 3,4, Late Aptian,
Krasnaya, SW Crimea, deposited in The Natural History Museum, London, no. PF 62921: dorsal and ventral
views (x21O). 5-7, Late Aptian, La Pena Formation, La Boca Canyon section, Mexico (from Longoria, 1974, pI.
16, figs 1-3): dorsal, peripheral and ventral views (xllO).
Figures 8-17 Lilliputianella bizonae (Chevalier, 1961). 8-10, earliest Late Aptian, La Bedoule, near Marseilles, SE France
(from Chevalier, 1962, pI. I, figs 24a-c), holotype deposited in the Laboratoire de Micropaleontologie, Universite
de Paris: ventral, peripheral and dorsal views (x85). 11-14, Late Aptian, sample N119, Kacha River, SW Crimea,
deposited in The Natural History Museum, London, nos PF 53085, PF 53086: 11, 12, dorsal and ventral views
(x115); 13, 14, ventral and dorsal views (xI20). 15-17, (=Hedbergella bollii Longoria, 1974), Late Aptian,
Schackoina cabri Zone, la Drome section, France (from Longoria, 1974, pI. 13, figs 12-14), holotype deposited
in the US National Museum of Natural History, Washington, DC, no. 184995: dorsal, peripheral and ventral views
(xI45).
Plate 9A

Figures 1-7 Wondersella athersuchi Banner and Strank, 1987. Late Aptian, highest Shuaiba Fonnation, Zakum-I Well, core
at 703 ft BRT, Abu Dhabi, United Arab Emirates, deposited in The Natural History Museum, London, thin-section
nos PF 52040 (holotype), PF 52041-PF 52043 (paratypes): 1-4, specimens showing development of angularity
and streptospirality, with umbilical closure well displayed in fig. 2 (holotype, fig. I) (x200); 5-7. reconstruction
of the external appearance of W. athersuchi (from Banner and Strank, 1987, p. 44, figs 3a--<:): 5, dorsal, ventral
and peripheral (adapertural) views (x200); fig. 6 shows the umbilicus incompletely closed, but a dashed line indi-
cates how the addition of one further chamber closes the umbilicus.
 10
The planispiral
Schackoinidae (Blowiella,
Claviblowiella gen.nov.,
Globigerinelloides,
Leupoldina and
Schackoina)
M.K. BouDagher-FadeI. F.T. Banner
and J.E. Whittaker

10.1 INTRODUCTION (which appeared in the Late Barremian, and had

bulbous chamber extensions) and the long-ranging
The planispiral immediate descendants of the Prae- Schackoina (which appeared in the Late Aptian,
hedbergellidae, the Schackoinidae, first evolved in ranged up into the Maastrichtian, and had elongate
the latest Early Barremian (Figure 10.1). The first slender tubulospines).
of this family were species of the genus Blowiella, The morphology of the different species of
a simple planispiral form, which had only four Blowiella, Globigerinelloides, and even Leupol-
chambers in the last whorl. We believe these to dina are regular and constant from individual
have arisen directly from Praehedbergella itself to individual. However, the genus Schackoina
(Figure 5.2). In the late Early Aptian, species of appears to be one of great morphological vari-
Blowiella evolved which had five chambers in the ability, even within the Early Cretaceous alone.
last planispiral whorl, and at almost the same time Many species names have been proposed to
these five-chambered Blowiella began to elongate accommodate these variations but we have only
their chambers radially; these high chambered considered the ones listed in this chapter to be
forms are grouped into the short-lived genus consistently recognizable. For example, some
Claviblowiella (gen.nov.) which became extinct at species possess only one tubulospine in the final
the close of the Aptian (BouDagher-Fadel, 1996) chamber but others may have two or three, or even
or soon after. Chambers became really high and more; we believe this to be the result of individual
peripherally elongate in the genera Leupoldina variation within one species, and as a result names
 ... ...
N
......
N 3:
~ ... D)
N

BARREMIAN APTIAN ALBIAN

EARLY LATE EARLY LATE EARLY LATE

Blowiella blowi s.s.

B. duboisi
B. gottisi

B. maridalensis

B. moulladei

B.solida

Claviblowiella mlnai

- c. saundersi
c. sigali
- Globigerinelloides algerianus
G. ferreolensis
Leupoldina protruberans
...
,. Schackoina cenomana

s. cabri
s. cepedai
...,. s. pentagonalis
Figure 10.1 Stratigraphical distribution of Early Cretaceous Blowiella, Claviblowiella and Leupoldina. Chronostratigraphy and Absolute Time Scale after Gradstein et al.
(1995)
 Systematics 179

like Schackoina multispina (= Hantkenina multi- Globigerinelloides at the beginning of the

spina Cushman and Wickenden, 1930) are not Globigerinelloides ferreolensis Zone of Late
conserved. Aptian time.
Globigerinelloides has a much greater number
of chambers than Blowiella, much broader
10.2 SYSTEMATICS umbilici and longer portici.

Superfamily GLOBIGERINACEA Carpenter, Range: Latest Early Barremian to Late Aptian.

Parker and Jones, 1862 This species (sensu stricto) was first described
from the 'early Middle' Aptian (the Leupoldina
Family SCHACKOINIDAE Pokorny, 1958 protuberans Zone) of the upper Cuche Formation
Genus Blowiella Kretchmar and Gorbachik, in Trinidad. Longoria (1974) illustrated this
1971, emend. Banner and Desai, 1988 species from the Late Aptian gorbachikae Zone
of La Perra, Mexico and the S. cabri Zone, of la
Type species Planomalina blowi Bolli, 1959 Drome, France. Butt's (1979) record of B. blowi
from DSDP Site 397 (sample 42-2) is of disputed
Blowiella blowi (Bolli, 1959) sensu stricto
age, being referred by Butt to the Barremian (?-
(Plate 10.1, Figs 1--4; Figure 10.1) Aptian) but being dated as Late Hauterivian by
Wind and Cepek (1979) on nannofloras. The work
1959 Planomalina blowi Bolli (pars), p. 260, published by Butt, at least, proves that the earliest
pI. 20, fig. 2a-b (holotype), (non fig. 3, record of Blowiella is older than Aptian. However,
paratype). the proposed phylogeny suggests to us that a Late
1974 Globigerinelloides blowi (Bolli), Longoria, Hauterivian age for any species of Blowiella is
p. 82, pI. 4, figs 11-13 (?4, ?7). very unlikely and that a Barremian age is much
1979 Globigerinelloides blowi (Bolli), Butt, more sustainable. Caron (1985) figured a specimen
p. 258, pI. 3, figs 14-17. from the Angola Basin, southeastern Atlantic
1985 Globigerinelloides blowi (Bolli), Caron, (thought to be 'reworked in Late Albian'). Ali-
p. 47, figs 29.10-11. Zade and Khalilov (1986) used B. blowi as a zonal
1995 Globigerinelloides blowi (Chevalier), index for the Early Aptian of Azerbaijan.
Weiss, pI. 1, figs 8,9. BouDagher-Fadel et al. (1997b) recorded this
1997b Blowiella blowi Bolli, BouDagher-Fadel et species in the Aptian of the Kacha River, SW
al., pI. 6, figs 1, 2. Crimea, and in the later part of the Early Aptian
of Tunisia. Finally, Weiss (1995) illustrated it
Remarks: Of the primary type series (Bolli, from the Early Aptian of the Wiechendorf 1/86
1959), only the holotype is characterized by Borehole, Lower Saxonian Basin, Germany.
having five (and a fraction) spherical chambers in This species has therefore a widespread distri-
the last whorl, increasing gradually in size. It bution, being found in both the northern margin
is biumbilicate, with a symmetrical aperture, of Tethys and in southern Tethys, as well as in
extending laterally into each umbilicus, furnished the Caribbean and the southeast Atlantic.
with portici. The four-chambered paratype of B.
blowi, with asymmetric final chambers is now Blowiella duboisi (Chevalier, 1961)
referred to B. moulladei (q.v.). (Plate 10.1, Figs 5-8; Figure 10.1)
Longoria (1974, plate 4) has illustrated the
increase in the chamber number in the descendants 1961 Globigerinella duboisi Chevalier, p. 33,
of the typical form of B. blowi s.s. to six to seven pI. 1, figs 14-18.
chambers in the final whorl. With an increase in 1974 cf. Globigerinelloides duboisi (Chevalier),
umbilical diameter, this can lead to the great Longoria, pp. 83, 84, pI. 4, figs 15, 16,
increase in chamber number which characterized pI. 11, figs 12, 13.
180 The planispiral Schackoinidae

1988 Blowiella duboisi (Chevalier), Banner and pp. 406-408, pI. 11, fig. 3 (topotype of
Desai, p. 172, p. 8, figs 10-12. Planomalina maridalensis Bolli).
1988 Blowiella maridalensis (Bolli), Banner and
Remarks: This species is characterized by its Desai, pp. 172, 173, pI. 9, figs 1-3.
high chambers (as high as long) and its excep- 1997b Blowiella maridalensis (Bolli), BouDagher-
tionally broad umbilicus (at least 25% of the Fadel et al., pI. 5, figs 1, 2.
greatest diameter).
Remarks: This species is distinguished by its
reniform chambers, which are distinctly longer
Range: Late Aptian. B. duboisi was first
than high, and its broad umbilicus.
described from the Gargasian of the south of
France, and was recorded from the Schackoina Range: Late Aptian. The primary types were
cabri Zone of Speeton, England by Banner and obtained by Bolli from the lower part of the
Desai (1988). Maridale Formation in Trinidad which spans from
Blowiella gottisi (Chevalier, 1961) the latest Aptian to the Early Albian. Longoria
(1974) illustrated this species from the Late Aptian
(Plate 10.1, Figs 9-12; Figure 10.1) cabri Zone of la Drome, France and from La Peiia,
1961 Globigerinella gottisi Chevalier, pp. 32, Mexico as well as from the ferreolensis Zone of
the stratotype Gargasian. Banner and Desai (1988)
33, pI. 1, figs 9a, b, pI. 1, figs 10-13.
1974 Globigerinelloides gOttlSI (Chevalier), recorded this species from the Schackoina cabri
Zone of Speeton Cliff, NE England, while
Longoria, p. 85, pI. 7, figs 10-13 (?7, 8).
BouDagher-Fadel et al. (1997b) found it from the
1988 Blowiella gottisi (Chevalier), Banner and
Late Aptian of the Kacha River, SW Crimea.
Desai, p. 172, pI. 8, figs 1-3.
1995 Globigerinelloides gottlSI (Chevalier), Blowiella moulladei BouDagher-Fadel, 1995
Weiss, pI. 1, figs 5, 6.
(Plate 10.2, Figs 1-3; Figure 10.1)
Remarks: This species distinguished from B. 1959 Planomalina blowi Bolli (pars), pI. 20, fig.
blowi primarily by its smaller umbilicus, and 3 (Paratype) (non holotype, figs 2a,b).
narrower, more restricted aperture. 1988 Blowiella saundersi (Bolli), Banner and
Desai, p. 173, pI. 9, figs 4-6.
Range: Aptian. Chevalier (1961) described his 1995 Blowiella moulladei BouDagher-Fadel,
primary types from the Gargasian (Late Aptian) p. 146, pI. 4, figs 1, 2.
of south France. Longoria (1974) has confirmed
its occurrence in the same stage of SE France and Remarks: This species is distinguished by
in the Schackoina cabri Zone of Mexico. Banner having asymmetrically high chambers, higher in
and Desai (1988) also obtained their specimens their posterior part. When the chambers become
from the cabri Zone at Speeton, while Weiss radially very elongated but still retain their char-
(1995) figured B. gottisi from the Early Aptian of acteristically asymmetric shape, Blowiella evolves
the Wiechendorf 1186 Borehole, Lower Saxony into the younger form Claviblowiella saundersi
Basin, Germany. (Bolli) (see p. 181 below).

Blowiella maridalensis (Bolli, 1959) Range: Latest Early Aptian-early Late Aptian.
(Plate 10.1, Figs 13-16; Figure 10.1) BouDagher-Fadel's 1995) figured specimens were
1959 Planomalina maridalensis Bolli, p. 261, pI. obtained from the latest part of the Early Aptian
20, figs 4-6. in Tunisia. Similar specimens were figured from
1974 GlobigerineUoides maridalensis (Bolli), the upper part of the S. cabri Zone of NE England
Longoria, p. 86, pI. 9, figs 4, 7,10-13. by Banner and Desai (1988) but were then incor-
1977 Globigerinelloides blowi (Bolli), Masters, rectly referred to B. saundersi.
 Systematics 181

Blowiella solida Kretchmar and Gorbachik, 1986 Claviblowiella saundersi (Bolli, 1959)
(Plate 10.2, Figs 4-6; Figure 10.1) (Plate 10.2, Figs 7-11, Plate 1OJ, Figs 6-9;
Figure 10.1)
1959 Planomalina escheri (Kaufmann), Bolli,
p. 260, pI. 20, figs 7, 8. 1959 Planomalina saundersi Bolli,p. 262, pI. 20,
1986 Blowiella solida Kretchmar and Gorbachik figs 9-11.
(in Gorbachik), p. 123, pI. 27, fig. 3. 1974 Globigerinelloides saundersi (Bolli), Long-
1997b Blowiella salida Kretchmar and Gorbachik; oria, p. 88, pI. 3, figs 2, 6-12, pI. 9, figs 8,9.
BouDagher-Fade1 et ai., pI. 6, figs 3,4. 1977 Globigerinelloides saundersi (Bolli),
Masters, p. 412, 413, pI. 11, fig. 6.
Remarks: This species has five chambers in the 1986 Blowiella saundersi (Bolli), Gorbachik,
last whorl, the first two subg10bu1ar, the last three pI. 27, fig. 4.
being broader than long. 1988 Blowiella saundersi (Bolli), Banner and
Desai, pp. 173, pI. 9, figs 4-6.
Range: Late Aptian. This species was originally 1995 Globigerinelloides saundersi (Bolli),
described from the Late Aptian of Crimea. It was Weiss, pI. 2, figs 1, 2.
reported, but mis-identified, as Planomalina
escheri (Kaufmann) by Bolli (1959) from the Late Remarks: This species is characterized by its
Aptian of Trinidad. BouDagher-Fade1 et al. found asymmetrically ovoid chambers, which are higher
it in the Late Aptian of Kacha River valley and in their posterior part than anteriorly. The speci-
of Krasnaya, SW Crimea. mens figured by Banner and Desai (1988, figs 4-6)
from the earliest part of the Late Aptian and those
Genus Claviblowiella gen.nav.
found by us in Jebel Chenanrafa, Tunisia, from
Type species: Blowiella saundersi Bolli, 1959 the latest part of the Early Aptian, show the phylo-
genetically early elongation of the chambers. This
Diagnosis: Claviblowiella is a genus of the radial elongation is more pronounced, highest in
Schackoinidae. The test is planispiral with four to mid-point, in the latest Aptian of the Maridale
six chambers in the last whorl, microperforate, Formation (Bolli, 1959, fig. 11, Masters, 1977, fig.
smooth, not muricate, but differs from Blowiella 6) and the Crimea (Gorbachik, 1986, fig. 4). The
in its chambers being radially elongate, higher specimens figured here from the Late Aptian of
than broad clavate. [Gender: feminine]. Crimea (Plate 10.2, Figs 7-11) show that the
ultimate and penultimate chambers can be asym-
Remarks: This tendency to radial elongation, metrically higher in their posterior part.
and the similarity in morphology exemplified by
the biumbilicate, four to six chambered planispire, Range: Latest Early Aptian to (possibly) earliest
smooth microperforate wall and forwardly Albian. The primary types were described from
projecting portici in the genus Claviblowiella is a the barri Zone of the Maridale Formation,
clear indication of its future evolution into Trinidad, which is believed to be latest Aptian to
Leupoldina (Banner and Desai, 1988, p. 173). Our earliest Albian in age (Bolli, 1959). It was
new genus differs from Leupoldina by the clavate recorded in the maridalensis and cabri zones,
breadth of its radially elongate chambers, lacking latest Early Aptian-early Late Aptian of SE France
the slender elongation typical of Leupoldina and and Mexico by Longoria (1974), the Wiechendorf
Schackoina. 1/86 Borehole, Lower Saxonian Basin, Germany
(Weiss, 1995) and at Speeton, NE England by
Range: There are three species assigned to Banner and Desai (1988). We also found it in the
Claviblowiella, and the genus appears to range Late Aptian of Krasnaya and the Kacha River
from the late Early Aptian to the end of the Aptian; section in SW Crimea and in the latest part of the
there is possibly an earliest Albian occurrence. Early Aptian of Tunisia.
182 The planispiral Schackoinidae

Claviblowiella miMi (Obregon de la Parra, 1959) Blowiella) in the Late Aptian. SEM photographs
(on Plates 10.4, 10.5) of paratypes of G. algeri-
(Plate 10.3, Figs 1-3; Figure 10.1)
anus (the type species) and topotypes of G. ferre-
1959 Globigerinelloides minai Obregon de la olensis (kindly lent by Dr M. Moullade, University
Parra, p. 151, pI. 4, figs 3a-c. of Lyon) show that this genus is also microper-
forate and non-muricate (although it commonly
Remarks: This species was described from develops perforation cones on the periphery of the
samples collected from the Aptian (Gargasian), La earliest chambers of the last whorl). However, it
Pefia Formation of Neuvo Laredo State, Mexico. has long been known that much larger, macro-
The original drawings were not good (refigured perforate homeomorphs existed (the two forms
here on Plate 10.3, Figs 1-3) but showed a micro- were contrasted by Banner and Desai, 1988, plate
perforate, smooth species with five chambers in 9, figs 7-9, but with uncertain nomenclature); they
the last whorl; the chambers of the later part of are now called Alanlordella BouDagher-Fadel
the last whorl were much higher than long or (1995), and are now known to be the earliest
broad, and the latest intercameral sutures were members of the Planomalinidae, the planispiral
deeply incised. The radial elongation of these analogues of the Hedbergellidae. These forms are
chambers indicate that the species must be referred described in detail in Chapter 12.
to Claviblowiella. We know of no subsequent
records of this species. Globigerinelloides algerianus Cushman and ten
Dam, 1948, sensu stricto
Range: Late Aptian of Mexico. (Plate 10.4, Figs 1-10; Figure 10.1)
Claviblowiella sigali (Longoria, 1974)
1948 Globigerinelloides algeriana Cushman and
(Plate 10.3, Figs 4, 5; Figure 10.1) ten Dam (sic), p. 43, pI. 8, figs 4-6.
1974 Globigerinelloides algerianus Cushman and
1974 Globigerinelloides sigali Longoria, p. 88,
ten Dam; Longoria, pp. 77-79, pI. 6, figs
pI. 3, fig. 1, pI. 7, figs 6,9. 1-18.
Remarks: This species has five chambers in the 1981 Globigerinelloides algeriana Cushman and
final whorl, increasing rapidly in size; the penulti- ten Dam (sic); Bellier and Chitta, p. 42,
mate chamber is as large as the first two cham- pI. 2, figs 16-18.
bers together, and the last one is very high. 1985 Globigerinelloides algeriana Cushman and
ten Dam (sic); Caron, p. 47, figs 29.5-7.
Range: Late Aptian. This species was first 1986 Globigerinelloides algerianus Cushman
described by Longoria (1974) from la Pefia and ten Dam; Gorbachik, p. 132, pI. 29,
Formation in Mexico. figs 2, 3.
1993 Globigerinelloides algerianus Cushman
Genus Globigerinelloides Cushman and ten and ten Dam; Shahin, p. 422, fig. 6.2.
Dam, 1948 1997b Globigerinelloides algerianus Cushman
and ten Dam; BouDagher-Fadel et al.,
Type species Globigerinelloides algerianus
pI. 4, figs 6, 7.
Cushman and ten Dam, 1948

Remarks: A marked increase in the number of Remarks: This species is distinguished by its
chambers in the last whorl (to seven or many completely planispiral and microperforate test
more), with a great widening of the umbilici, which lacks a keel; by its high number of cham-
lengthening of the portici (which become draped bers, ten to twelve in the last formed whorl; and
backwards from almost all chambers of the last by an equatorial aperture with relict openings in
whorl) and lateral compression of the test, gave broad umbilici and long portici on both sides of
rise to Globigerinelloides (from ancestral the test.
 Systematics 183

The stratigraphic range of G. algerianus sensu 1961 Biticinella ferreolensis Moullade, p. 4, pI.
stricto overlaps in the Late Aptian of Tunisia and 1, figs 1-5.
Kacha River with the older taxon Globigerinoides 1966 Globigerinelloides ferreolensis (Moullade),
ferreolensis (Moullade). According to Prof. M. Moullade, p. 123, pI. 9, figs 1-3.
Caron (personal communication, 1995) the only 1974 Globigerinelloides ferreolensis (Moullade),
difference between the younger form G. algerianus Longoria, pp. 84, pI. 5, figs 7,8, pI. 8, figs
and the older form G. ferreolensis is the greater 1-3,8-15, pI. 14, figs 7, 8, pI. 27, figs 3,4,
number of chambers per whorl in the former. 12.
However, the evolution is demonstrated by the 1974 Globigerinelloides macrocameratus Long-
consistent, gradual change in the morphological oria, pp. 85,86, pI. 5, figs 1-6.
grade from G. ferreolensis (Moullade) into G. 1981 Globigerinelloides ferreolensis (Moullade),
algerianus s.s. seen in our Crimean and Tunisian Bellier and Chitta, p. 42, pI. 2, figs 13-15.
material. Their constant presence in the same 1986 Globigerinelloides ferreolensis (Moullade),
samples in Crimea and Tunisia agrees with the Gorbachik, pI. 29, fig.I.
coincidence in ranges of the two taxa in the middle 1987 Globigerinelloides ferreolensis (Moullade),
part of the Late Aptian (algerianus Zone of Bellier Ben Haj Ali, p. 83, pI. 1, figs 1,2.
and Chitta, 1981; Caron, 1985; Ben Haj Ali, 1987; 1991 Globigerinelloides ferreolensis (Moullade),
and Altiner, 1991). Altiner, pI. 16, fig. 1.
Globigerinelloides arose rapidly from the 1993 Globigerinelloidesferreolensis (Moullade),
equally planispiral Blowiella by the addition of Shahin, p. 430, pI. 6, fig. 2.
more chambers and broader umbilici and longer 1995 Globigerenelloidesferreolensis (Moullade),
portici to give rise to G. ferreolensis and algeri- BouDagher-Fadel, p. 147, pI. 2, figs 8,9.
anus which in tum gave rise to Planomalina
cheniourensis (Sigal) (Caron, 1985) and to P. Remarks: This species differs from the strati-
buxtorji (Gandolfi), via Alanlordella, by acquiring graphically younger taxon G. algerianus Cushman
a macroperforate test and, progressively, a and ten Dam only in the smaller number of
peripheral keel. For further discussions, see chambers in the last whorl. While G. ferreolensis
Chapter 12. has eight chambers in the last-formed whorl, G.
algerianus has a larger number of chambers, ten
Range: G. algerianus s.s. occurs within the Late to twelve in the same whorl. The gradual morpho-
Aptian. According to Caron (1985), it ranges from logical change between the two forms is consis-
the top of the Schackoina cabri Zone to the base tent throughout the samples in Crimea and
of the Blefuscuiana gorbachikae Zone. It was orig- Tunisia.
inally described by Cushman and ten Dam (1948) The species arises rapidly from the equally
from the Late Aptian of western Algeria. It was planispiral but much less compressed form
reported from the Late Aptian of France, Mexico Blowiella blowi (Bolli) which has smaller umbilici
(Longoria, 1974), Tunisia (Bellier and Chitta, and even fewer chambers in each whorl.
1981; Ben Haj Ali, 1987), the Crimea (Gorbachik, The new species described by Longoria (1974)
1986; BouDagher-Fadel et al., 1997b), Tunisia as G. macrocameratus is very similar to G.
(BouDagher-Fadel, 1995, fig. 1) and from Turkey ferreolensis and should be considered as its
(Shahin, 1991). It has not been found in the North junior synonym; it is refigured on Plate 10.5, Figs
Sea area either from offshore and in the Aptian 13-15.
outcrops at Speeton, coast of NE England. The
stratigraphically older species G. ferreolensis Range: Late Aptian. G. ferreolensis was first
(Moullade) has a longer range (see below). described from the Early Gargasian (earliest Late
Aptian) of la Drome, France. It was also found in
Globigerinelloides ferreolensis (Moullade, 1961)
the Late Aptian of France and Mexico (Longoria,
(Plate 10.5, Figs 1-15; Figure 10.1) 1974), Tunisia (Bellier and Chitta, 1981; Ben Haj
184 The pJanispiral Schackoinidae

Ali, 1987; BouDagher-Fadel, 1995), the Crimea 1997b Leupoldina protuberans Bolli; BouDagher-
(Gorbachik, 1986; BouDagher-Fadel et al., 1997b), Fadel et al., pI. 5, figs 8, 9.
Turkey (Altiner, 1991) and Egypt (Shahin, 1993).
Genus Leupoldina Bolli, 1957 emend. Banner Remarks: This form consists of four to five
and Desai, 1988 chambers in the last whorl, each elongated with a
subglobular extension. These extensions are
Type species Leupoldina protuberans Bolli,
mostly broken. In the adult the last whorl will
1957
remain planispiral but the last few chambers may
Leupoldina protuberans Bolli, 1957 become biserial or triseriaI.
(Plate 10.6, Figs 1-7; Figure 10.1) In the original publication (Bolli, 1957), L.
1957 Leupoldina protuberans Bolli, p. 277, pI. protuberans was (otherwise beautifully) drawn (by
2, figs 1-13. L. Isham) with macroperforations; this is an
1957 Schackoina pustulans pustulans Bolli, example of the artist drawing what he presumed
p. 274, pI. 1, figs 1-7. to exist, even though he has not seen it. L. protu-
1957 Schackoina pustulans quinquecamerata berans, like all other species of Leupoldina, is a
Bolli, p. 275, pI. 1, figs 1-7. smooth microperforate member of the Schacko-
1957 Schackoina reicheli Bolli, p. 275, pI. 1, figs inidae. The various forms of preservation of the
8-10. adult test have given rise to the plethora of junior
1959 Schackoina reicheli Bolli; Sigal, p. 74, figs synonyms listed above. Breakage of the bulbous
48,49. terminations of the tubulospines (which is the
1971 Leupoldina reicheli (Bolli); Gorbachik, main criterion for differentiation between Leupol-
p. 138, pI. 30, figs 8a, b. dina and Schackoina) often makes precise identi-
1974 Leupoldina reicheli (Bolli), Longoria, p. 92, fication somewhat difficult.
pI. 27, fig. 17.
1977 Leupoldina pentagonalis (Reichel), Mas- Range: Latest Barremian to Early Albian.
ters, pp. 425, 426, pI. 14, figs 2, 3. Leupoldina protuberans (and its synonyms) was
1984 Leupoldina pustulans (Bolli), Salaj, pp. first described from the Early Cretaceous (Aptian
856, 857, pI. 3, fig. 10. to Albian) of Trinidad. This species was also found
1985 Schackoina cabri Sigal; Caron, figs 35. in the Late Aptian of the Kacha River in the Crimea
2-4. (Gorbachik, 1986; BouDagher-Fadel et al., 1997b),
1986 Leupoldina protuberans Bolli; Gorbachik, the Wiechendorf 1186 Borehole of Germany
pI. 28, fig. 1. (Weiss, 1995), latest Barremian to Early Aptian
1986 Leupoldina pustulans (Bolli), Gorbachik, (Salaj, 1984) and early Late Aptian (Early
pI. 28, fig. 2. 'Gargasian') (Ben Haj Ali, 1987) and latest Early
1986 Leupoldina reicheli (Bolli), Gorbachik, Aptian to Late Aptian (BouDagher-Fadel, 1995) of
pI. 28, fig. 3. Tunisia.
1987 Leupoldina cabri (Sigal), Ben Haj Ali,
p. 83, pI. 3, figs 4-10. Genus Schackoina Thalmann, 1932
1987 Schackoina pustulans Bolli; Ben Haj Ali, Type species Siderolina cenomana Schacko,
pI. 3, figs 11, 12. 1897
1995 Leupoldina protuberans Bolli; BouDagher-
Fadel, p. 148, pI. 4, figs 3-5. Schackoina cenomana (Schacko, 1897)
1995 Leupoldina gr. cabri (Sigal), BouDagher-
(Plate 10.7, Figs 1-5,7; Figure 10.1)
Fadel, p. 147, pI. 4, figs 8, 9.
1995 Leupoldina cabri (Sigal), Weiss, pI. 2, figs 1897 Siderolina cenomana Schacko, p. 166,
6-8. text-figs 4 (3-5).
1995 Leupoldina pustulans (Bolli), Weiss, pI. 2, 1932 Schackoina cenomana (Schacko), Thal-
figs 9-12. mann, p. 289.
 Systematics 185

1957 Schackoina cenomana (Schacko), Bolli, tions to the elongate chambers typical of Leupol-
Loeblich and Tappan, p. 26, pI. 2, figs 1, dina, even though, like Leupoldina, its last cham-
2 (near topotypes). bers may become bi- or even tri-serial, with its
1960 Schackoina cenomana cenomana tubulospines becoming multiple on the final cham-
(Schacko), Masella, pp. 11-13, text-figs bers. Schackoina differs from the Eocene genus
1(1-9), 2(1-4), 10(1-4) (q.v. for syn- Hantkenina, its larger, gross homeomorph, in
onymy). possessing the smooth, microperforate wall typical
of the Schackoinidae and Praehedbergellidae, and
Remarks: In 1897, Schacko described random also in possessing trochospiral initial whorls and
thin-sections of planktonic foraminifera from the in having tubulospines which are thin-walled yet
Cenomanian Chalk of Moltzow, Mecklenburg, having broad inner spaces (Hantkenina has tubu-
Germany; these specimens had four chambers in lospines which are thick-walled with very narrow
the last of a succession of whorls, and these late inner spaces); these two genera are thus easily
chambers were narrowly elongated into 'tubulo- distinguished both in the solid specimen and in
spines' (sensu Bolli, Loeblich and Tappan, 1957). random thin-section.
Schacko (1897) called these forms Siderolina Schackoina cenomana has four chambers in the
cenomana, but Siderolina is a d'Orbigny genus last whorl and is very involute; its umbilici are
(recte Siderolites) which is totally unrelated; very small and narrow.
Thalmann (1932) placed them in his new genus
Schackoina as its type species. The existence of Range: Late Aptian to Cenomanian (it may even
the type specimens of S. cenomana has never been extend into the Coniacian). This species was first
established but hypotypes ('near topotypes') were described from the Cenomanian of Germany. We
figured by Bolli, Loeblich and Tappan (1957, pI. have found it and illustrate it here from the Albian
2, figs 1, 2) from the Cenomanian of Luneburg, of Tunisia.
Germany. One of these specimens (Ibid., plate 2,
figs la-c; reproduced in our Plate 10.7, Figs 1-3) Schackoina cabri S'igal, 1952
is formally chosen here as the neotype, because it (Plate 10.6, Figs 8-14; Figure 10.1)
matches the original syntypes and is from a
locality as near as possible to them and is, as far 1952 Schackoina cabri Sigal, pp. 20,21, fig. 18.
as we know, of the same age. Masella (1960) 1985 Schackoina cabri Sigal; Caron, figs 35.1-4.
figured very many specimens of Schackoina (solid 1981 Schackoina cabri Sigal; Bellier and Chitta,
and thin-sectioned) from the Cenomanian of p. 42, pI. 3, figs 22-28.
Sicily; she referred them to many subspecies 1988 Leupoldina gr. cabri (Sigal), Banner and
of cenomana, including four-chambered forms as Desai, p. 176, pI. 10, figs 1-8.
S. cenomana s .s. (believed to range from the
Cenomanian to the Turonian), five-chambered Remarks: This species is evolute and is char-
forms as S. pentagonalis Reichel, 1948 (Albian- acterized by having four to five chambers in the
Coniacian) and many others. Many of the thin last whorl, which develop tubulospines.
sections drawn by Masella (1960) showed initial S. cabri is characterized by its broad, evolute
trochospiral whorls (buried by the adult involute umbilici. It may become bispinate in its last cham-
planispiral) and the tubulospines of the early bers, but it may remain unispinate (one spine on
planispiral whorls preserved inside the adult cham- each chamber) throughout.
bers (which usually have their spines broken oft).
The youngest Schackoina was recorded from the Range: Late Aptian to Early Albian. Schackoina
Maastrichtian by Aurouze and de Klasz (1954). cabri is cosmopolitan. It was first described from
Schackoina differs from the older and shorter- the Aptian of Jebel Rhazouane in Tunisia. It has
ranged Leupoldina in having narrowly elongate been reported from Trinidad, the Haute Alpes
tubulospines and in lacking the bulbous termina- (Longoria, 1974), Mexico (Caron, 1985), Speeton,
186 The planispiral Schackoinidae

NE England (Banner and Desai, 1988, pp. Blowiella, a genus which became extinct at the
173-174), and Tunisia (Bellier and Chitta, 1981). end of the Aptian. Four lines of evolution derived
new genera from this ancestor (Figure 5.2). The
Schackoina cepedai (Obregon de la Parra, 1959)
first was Leupoldina which appeared at the end of
(Plate 10.7, Figs 10-12; Figure 10.1) the Barremian by developing radially elongate
chambers with bulbous terminations; this genus
1959 Hastigerinelloides cepedai Obregon de la
persisted until the Early Albian, but in the Late
Parra, p. 151, pI. 4, fig. 5.
Aptian gave rise to Schackoina with slender
1974 Globigerinelloides cepedai (Obregon),
tubulospines. Schackoina was unique in the
Longoria, p. 83, pI. 7, figs 1-5, pI. 9, figs
Schackoinidae by persisting through the Late
1-3.
Cretaceous until the Maastrichtian. A much
shorter-lived genus was Claviblowiella gen.nov.,
Remarks: This species has five to six chambers
which evolved directly from Blowiella by forming
in the last whorl. The tubulospines are developed
broad elongations to its last chambers; this taxon
on only the last two or three chambers of the last
is known only from the highest Early Aptian to
whorl; the earlier chambers of the last whorl are
the latest Aptian (?earliest Albian). The fourth
subglobular. The umbilici are moderately broad.
lineage was one which became very important
biostratigraphically; the simple planispiral whorls
Range: Late Aptian of Mexico and Spain. became many-chambered in the Late Aptian, and
Schackoina pentagonalis Reichel, 1948 the umbilici broadened to form Globigeri-
nelloides; in the latest Aptian, this genus gave rise
(Plate 10.7, Figs 6, 8, 9; Figure 10.1) to the macroperforate (later muricate) Plano-
1948 Schackoina pentagonalis Reichel, p. 395, malinidae, which now provide the biostratigrapher
text-fig. l(a-f), 6(1), 7(1). with some easily identified, very distinctive
1960 Schackoina cenomana pentagonalis markers for the zonation of Albian deep-water
(Reichel), Masella, pp. 13, 14, text-figs sediments.
4(5, 9-14), 5(1-5). As we discuss further in Chapter 13, the radial
1995 Schackoina reicheli Bolli; BouDagher- elongation of the chambers in Claviblowiella,
Fadel, pI. 4, fig. 10. Leupoldina and Schackoina was probably due to
their presence in seawater undersaturated in
Remarks: This species has five chambers in the oxygen; this could explain the sporadic and often
last whorl, with a moderately sized and a partly rare occurrence of these fossil taxa. In contrast,
involute umbilicus. the Schackoinidae, which did not radially elongate
their chambers, were widespread in the palaeo-
Range: Late Aptian-Cenomanian. tropics (now intercontinental occurrence), and are
valuable biostratigraphic indices worldwide.
There are certain areas (e.g. the central North
10.3 CONCLUDING REMARKS Sea areas) where the planispiral Schackoinidae
have not yet been found. This might be due to the
The planispiral, microperforate, nonmuricate taxa population of these areas being derived principally
which we group into the family Schackoinidae first from the colder source areas of the northerly
appeared in the Early Barremian with the first climate.
Plate 10.1

Figures 1-4 Blowiella blowi (Bolli. 1959) sensu stricto. 1, 2, Late Aptian, Cuche Fonnation, Trinidad (from Bolli, 1959,
pI. 20, figs 2a, b), holotype deposited in the US National Museum of Natural History, Washington, DC, no.
263603: side and peripheral views (xl 10). 3,4, Early Aptian, sample N119, Kacha River, SW Crimea, deposited
in The Natural History Museum, London, no. PF 53082: side views, showing chambers gradually increasing in
size (x305).
Figures 5-8 Blowiella duboisi (Chevalier, 1961). 5, 6, Aptian (Gargasian), Pierre Fiche, western Provence, France (from
Chevalier, 1961, pI. 1, figs 14a, b), holotype deposited in the Laboratoire de Micropaleontologie, Universite de
Paris: side and peripheral views (x85). 7,8, earliest Late Aptian, Schackoina cabri Zone, sample 8109, Speeton
Cliff, NE England, deposited in The Natural History Museum, London, nos PF 52151, PF 52152: side views,
with high chambers and a broad umbilicus (x215 and x230, respectively).
Figures 9-12 Blowiella gottisi (Chevalier, 1961). 9, 10, Aptian (Gargasian), Pierre Fiche, western Provence, France (from
Chevalier, 1961, pI. I, figs 9a, b), holotype deposited in the Laboratoire de Micropaleontologie, Universite de
Paris: side and peripheral views (x80). 11, 12, earliest Late Aptian, Schackoina cabri Zone, sample 8109,
Speeton Cliff, deposited in The Natural History Museum, London, nos PF 52154, PF 52155: peripheral view
(x 150) and side view showing a small umbilicns (x 185).
Figures 13-16 Blowiella maridalensis (Bolli, 1959). 13, Late Aptian, lower Maridale Fonnation, Trinidad (from Bolli, 1959,
pI. 20, fig. 6a), holotype deposited in the US National Museum of Natural History, Washington, DC, no. 626329:
side view (xllO). 14, early Late Aptian, sample 8108, Schackoina cabri Zone, Speeton Cliff, NE England,
deposited in The Natural History Museum, London, no. PF 52142: side view (xI55). IS, 16, Late Aptian, sample
N120, Kacha River, SW Crimea, deposited in The Natural History Museum, London, no. PF 53086: side views
showing chambers longer than high and a broad umbilicus (x 165).
Plate 10.2

Figures 1-3 Blowiella moulladei BouDagher-Fadel, 1995. 1,2, early Aptian, SS356, Jebel Chenanrafa, holotype deposited in
The Natural History Museum, London, no. PF 53141: side views with chambers higher in their posterior part
than anteriorly (xI95). 3, Late Aptian, upper Schackoina cabri Zone, sample 8106, Speeton Cliff, NE England
(figured by Banner and Desai, 1988, pI. 9, fig. 4 as Blowiella saundersi (Bolli, 1959)), deposited in The Natural
History Museum, London, no. PF 52145: side view (xI90).
Figures 4-6 Blowiella solida Kretchmar and GorbachiJc, 1986, Late Aptian, Krasnaya, SW Crimea (from Gorbachik, 1986,
pI. 27, fig. 3), topotype deposited in the Geological Faculty of Moscow State University, Russia: side view (xI70).
5, 6, topotype, same locality, deposited in The Natural History Museum, London, no. PF 62920: side views
showing broad and shallow umbilici (x 170).
Figures 7-11 Claviblowiella saundersi (Bolli, 1959). Late Aptian, Krasnaya, SW Crimea, deposited in The Natural History
Museum, London, nos PF 62921, PF 62922: 7, 8, 10, 11, side views of respective specimens (x 105 and x 135);
9, enlargement of the asymmetrically ovoid last chamber showing the microperforations (x21O).
Plate 10.3

Figures 1-3 Claviblowiella minai (Obreg6n de la Parra, 1959). Late Aptian, La Pefia Formation, Mexico (from Obreg6n de la
Parra, 1959, pI. 4, figs 3a-c), holotype (depository not given): 1,3, side views; 2, peripheral view (x80).
Figures 4-5 Claviblowiella sigali (Longoria, 1974). Late Aptian, Schackoina cabri Zone, La Pefia Formation, La Boca Canyon,
Mexico (from Longoria, 1974, pI. 7, figs 6, 9), holotype deposited in the US National Museum of Natural History,
Washington, DC, no. 185024: side view (with a very high last chamber), and a peripheral view. (Dimensions not
given.)
Figures 6-9 Claviblowiella saundersi (Bolli, 1959). Latest Aptian (- ?earliest Albian), Maridale Formation, Trinidad (from
Bolli, 1959, pI. 20, figs 9-11), deposited in the US National Museum of Natural History, Washington, DC, nos
626307 (holotype), 626305, 626306 (paratypes): 6,9, side views of two paratypes (x95); 7, 8, peripheral and side
views of the holotype (x95).
Plate lOA

Figures 1-11 Globigerinelloides algerianus Cushman and ten Dam, 1948, sensu stricto. 1,2, Late Aptian. Jebel Menaouer, near
Relizane, W Algeria (from Cushman and ten Dam, 1948, pI. 8, figs 5, 6): I, holotype (side view, x65); 2, paratype
(peripheral view, x70); 3-9, paratype deposited in the US National Museum of Natural History, Washington, DC,
56790a: 3, peripheral view showing a completely planispiral test; 4,5, side views (x70), 8, enlargement of fig.
4 (xI25); 6--9, enlargement of the wall surface showing microperforations (fig. 7, x500) and perforation cones
(figs 6 and 9) (x420 and x300, respectively). 10, II, Late Aptian, Krasnaya, SW Crimea, deposited in The Natural
History Museum, London, nos PF 62909, PF 62910: side views showing broad umbilici and long portiei (x110).
Plate 10.5

Figures 1-15 Globigerinelloides ferreolensis (Moullade, 1961). 1--6, Late Aptian (Gargasian), Saint-Ferreol-Trente-Pas, la
Dr6me, SE France (from Moullade, 1961, pI. I, figs 1-5) types deposited in the Department of Earth Sciences,
University of Lyon, France: 1,3,4, paratype, side, peripheral and (other) side view (x60); 2,5, holotype, side
views (x60); 6-10, two topotypes kindly leant by Prof. Moullade and photographed by SEM at The Natural
History Museum, London: 6,7, side views (xI40); 8,9, side views of other specimen (x1l5), 10, enlargement
of the wall surface of the early whorls, showing perforation cones (xllDO). II, 12, Late Aptian, SS534, Beauvoir-
IV Well, Tunisia, deposited in The Natural History Museum, London, nos PF 62907, PF 62908: 11, side view
showing a microperforate test probably with perforation cones on the early chambers of the last whorl (x135);
12, enlargement of wall showing microperforations (x3200). 13-15, (= Globigerinelloides macrocameratus
Longoria, 1974) (from Longoria, 1974, pI. 5, figs 2-4), holotype deposited in the US National Museum of Natural
History, Washington, DC, no. 185022: side, peripheral and side views (x 100).
Plate 10.6

Figures 1-7 Leupoldina protuberans Bolli, 1957. 1,2, probable Aptian, Piparo River, Trinidad, holotype (from Bolli, 1957,
pI. 2, figs 7, 7a), deposited in the Natural History Museum, Basle, Switzerland, no. C 2437: side and peripheral
views (xl40). 3,4, Late Aptian, Krasnaya, SW Crimea, deposited in The Natural History Museum, London, no.
PF 62911: side views with broken subglobular extensions (3, x130; 4, xI25). 5, earliest Late Aptian, Schackoina
cabri Zone, sample 8109, Speeton Cliff, NE England, (previously figured by Banner and Desai, 1988, pI. 10, fig.
5, as Leupo/dina gr. cabri (Sigal)), deposited in The Natural History Museum, London, no. PF 521336: side view
(xl90). 6, 7, Late Aptian, sample SS389, Jebel Chenanrafa, Tunisia (previously figured as Leupoldina gr. cabri
(Sigal) by BouDagher-Fadel (1995, pI. 4, fig. 8)), deposited in The Natural History Museum, London, no. PF
53146: side views (xI20).
Figures 8-14 Schackoina cabri Sigal, 1952. 8,9, Aptian, Jebel Rhazouane, Tunisia (from Sigal, 1952, p. 21, text-fig. 18), holo-
type deposited in the author's personal collection, no. 101: side and peripheral view (x90). 10-14, early Late
Aptian, Schackoina cabri Zone, sample 8109, Speeton Cliff, NE England (previously figured by Banner and Desai,
1988, pI. 10, fig. 5, as Leupo/dina gr. cabri (Sigal)), deposited in The Natural History Museum, London, nos PF
52132-PF 52135: side views showing chambers developing long tubulospines; 10, with bifurcation of chamber
extension (10, x195; 11, x200; 12, x340; 13, x180; 14, x350).
Plate 10.7

Figures 1-5,7 Schackoina cenomana (Schacko, 1897). 1-3, Cenomanian, Germany, hypotype (here formally designated
neotype) deposited in the US National Museum of Natural History, Washington, DC, no. P 4644b: 1,3, oppo-
site sides showing radially elongate chambers and basal portion of the tubulospines; 2, peripheral view showing
low equatorial aperture (x210). 4, 5, 7, Albian, sample SS534, Beauvoir-IV Well, Tunisia, deposited in The
Natural History Museum, London, nos PF 62927, PF 62928; side views (4, x155; 5, x95; 7, x135).
Figures 6, 8,9 Schackoina pentagonalis Reichel, 1948.6, Late Aptian, sample SS577, Jebel Oust-III Well (previously figured
by BouDagher-Fadel, 1995, pI. 4, fig. 10 as Schackoina reicheli (Bolli)), deposited in The Natural History
Museum, London, no. PF 53148: side view (xI15). 8,9, Late Aptian, sample SS534, Beauvoir-IV Well, deposited
in The Natural History Museum, London, nos PF 62927, PF 62928; side views showing a partly involute
umbilicus (8, xll0; 9, x115).
Figures 10--12 Schackoina cepedai (Obregon de la Parra, 1959). 10, Aptian (Gargasian), La Peiia Formation, Mexico, holo-
type (from Obregon de la Parra, 1959, pI. 4, fig. 5), depository not given: side view (xI50). 11, 12, Aptian,
Argos Formation, Spain (from Longoria, 1974, pI. 7, figs 1,2): side views (xI60).
 11
The evolution of
the Hedbergellidae
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

11.1 INTRODUCTION while the ancestor, B. praetrocoidea, is known

only from eastern Tethyan areas.
Around latest Early Aptian times (Figure 5.2) small
microperforate Blefuscuiana appear to have begun
colonizing deeper waters, thus evolving into larger 11.2 SYSTEMATICS
and macroperforate Hedbergella (BouDagher-
Fadel, 1996). Superfamily GLOBIGERINACEA Carpenter,
An example of this evolution is described in Parker and Jones, 1862
BouDagher-Fadel et al. (l997a): Hedbergella
trocoidea (Gandolfi) evolves from Blefuscuiana Family HEDBERGELLIDAE Loeblich and
praetrocoidea (Kretchmar and Gorbachik) in the Tappan, 1961
Aptian. Hedbergella trocoidea (Late Aptian to
earliest Late Albian) was the immediate ancestor Genus Hedbergella Bronnimann and Brown,
of Ticinella roberti (Gandolfi) (latest Aptian to the 1958
end of the Albian), the type species of Ticinella
Type species Anomalina lorneiana d'Orbigny
Reichel (Figure 8.2). This evolution is marked by
var. trocoidea Gandolfi, 1942
a gradual increase of the size of the test, an
increase in the number of chambers in the whorl, Hedbergella trocoidea (Gandolfi, 1942)
an evolution of macroperforations and muricae in
(Plate Ll, Fig. 7, Plate ILl, Fig. 4; Figure 8.2)
H. trocoidea, a widening of the umbilicus which
reaches its maximum in T. roberti where the 1942 Anomalina lorneiana d'Orbigny var.
portici are distally fused forming umbilical acces- trocoidea Gandolfi, p. 99, pI. 2, figs la-c,
sory apertures. Concomitant with this evolution is pI. 4, figs 2, 3, pI. 3, figs 2a, b, Sa, b.
an increase in palaeogeographic distribution, so 1974 Hedbergella trocoidea (Gondolfi), Long-
that T. roberti and H. trocoidea are worldwide, oria, p. 69, pI. 17, figs 1-16, pI. 18, figs 3-5.
204 The evolution of the Hedbergellidae

1986 Hedbergella trocoidea (Gandolfi), 11.3 CONCLUDING COMMENTS

Gorbachik, pp. 93-95, pI. 17, figs 1-5.
1987 Hedbergella trocoidea (Gandolfi), Ben Haj The macroperforations which characterize the
Ali, p. 83, pI. 2, figs 1-3,9-11. Hedbergellidae soon were accompanied by the
1991 Hedbergella trocoidea (Gandolfi), Altiner, development of muricae, from which murico-
p. 170, pI. 15, figs 7-12. carinae were later to develop (but never seen in
1993 Hedbergella trochoidea (Gandolfi) (sic), the Praehedbergellidae). In the latest Aptian, the
Shahin, p. 423. first Rotaliporidae appeared, their macroperforate
1995 Hedbergella trocoidea (Gandolfi), and muricate tests reflecting their ancestry in
BouDagher-Fadel,pp.150,151,pI.2,fig.13. Hedbergella (e.g. H. planisira (Tappan) (Plate
1997a Hedbergella trocoidea (Gandolfi), 11.1, Figs 1-3 but now developing accessory
BouDagher-Fadel et al., p.5, pI. 2, figs 1-9 umbilical apertures which also were never seen in
(q.v. for full synonymy). the Praehedbergellidae.
The muricae could fuse around the periphery of
Remarks: This species is characterized by the test to form a muricocarina. These early
having six to seven chambers in the last whorl, a muricocarinate members of the Hedbergellidae are
convex dorsal side and a concave ventral side with called Praeglobotruncana; some examples are
a broad umbilicus. It evolved from Blefuscuiana shown in Plate 11.1 (P. delrioensis (Plummer);
praetrocoidea (Kretchmar and Gorbachik) in the figs 5-7 and P. stephani (Gandolfi); figs 8-10).
Aptian by acquiring muricae over the early whorls A similar peripheral muricocarina was devel-
and macroperforations (BouDagher-Fadel et al., oped in the Planomalinidae (Chapter 12) but in
1997a). this family the portici in the broad umbilici always
remained separate. In contrast, the portici of
Range: Late Aptian-early Late Albian. This Praeglobotruncana changed, in the later evolution
species was originally described from the Breggia through Thalmanninella, into the Cenomanian
River section of Switzerland by Gandolfi (1942). Rotalipora, when the portici partially fused
Unlike its immediate ancestor, B. praetrocoidea, leaving accessory apertures open in the ventral
it has a cosmopolitan distribution. It has been sutures. Different fusion of the portici, and devel-
reported from the probable Lower Cretaceous of opment of further muricocarinae, led to the
California (Cushman and Todd, 1948, pI. 16, figs development of the very many distinctive genera
18, 19, as Globigerina almadenensis); the Albian of the Globotruncanidae. All these evolved from
of the Maridale Formation of Trinidad (Bolli, ancestral species of HedbergeUa which itself
1959, pI. 22, figs 5-7, as Praeglobotruncana rohri; originated, as we have seen, from Blefuscuiana of
the Albian of Romania (Neagu, 1965, pI. 10, figs the Early Cretaceous.
1, 2, 4, as Hedbergella planispira); from the It has been postulated (Banner and Desai, 1988)
upper Gargasian to 'Middle' Albian of France that the small, microperforate, smooth species of
(Moullade, 1966); from the uppermost Aptian to Belfuscuiana lived very near the surface of the
'Middle' Albian of the Bavarian Calcareous Alps ocean, and that their evolution into the larger,
(Risch, 1971); from the Late Aptian to the Early macroperforate, muricate species of Hedbergella
Albian in northern Mexico (Longoria, 1974); from (Plate 11.1, Figs 1-4) was accompanied by an
the Late Aptian to earliest Albian (Caron, 1985); adaptation to feeding in deeper, subsurface waters.
from the Aptian and Lower Albian of Tunisia The evolution of the geographically restricted B.
(Salaj, 1980; Bellier and Chitta, 1981; Ben Haj praetrocoidea (Kretchmar and Gorbachik) (plate
Ali, 1987; BouDagher-Fadel, 1995); from the 8.9, Figs 4-12) to the almost cosmopolitan H. tro-
Late Aptian into the Early Albian of Crimea coidea (Gandolfi) (plate 11.1, Fig. 4) indicates the
(Gorbachik, 1986; BouDagher-Fadel et al. 1997a); habitation of deeper environments in the Late
from the Late Aptian of Turkey (Altiner, 1991), Aptian-Early Albian, which allowed Hedbergella to
and northern Sinai, Egypt (Shahin, 1993). become much more widespread and cosmopolitan.
 Concluding comments 205

It may be of interest that Haq et al. (1988) indi- being responsible for the turnover in the plank-
cate that the Aptian-Early Albian was a time of tonic foraminiferal record.
significant eustatic sea-level change. According to Alexander (1985 MS) studied the cytoplasm and
their chart, major sea-level rises occur at the base pseudopodia of many living benthic foraminifera
of the Schackoina cabri Zone (early Late Aptian) and noted that the umbilicus became an extrathala-
and at the Aptian!Albian boundary. An overall mous digestive zone for ingested nutrient particles;
pattern of moderately falling, eustatic sea-level in he also noted that the characters of the umbilical
the Aptian was replaced by a pattern of rising sea- cytoplasm differed from those of the intrathalamous
level in the Albian, initiated by the transgressive cytoplasm and the pseudopodia, and that the latter
event at the Aptian!Albian boundary. This even- had a sharper differentiation when the umbilicus
tually culminated in the mid-Cretaceous sea-level was partly covered by the skeletal structure of broad
maxima. The rising sea-level pattern in the Albian aperturallips or umbilical plates. Banner and Desai
described by Haq et al. (1988) may correspond (1988) correlated these findings with the develop-
with the spread of deeper environments in what is ment of umbilical structures in the Globigerinacea,
now Europe, created by the broadening and deep- and from this it can be suggested that the broaden-
ening of the Proto-Atlantic. We postulate that this ing of the portici and their subsequent fusion in the
would account for the wide distribution of H. Hedbergella trocoidea-Ticinella roberti evolution-
trocoidea. The radiation and geographic dispersal ary lineage results from a skeletally more concealed
of true Hedbergella may also be related to the and bigger digestive zone in the organism's cyto-
cessation of the Aptian oceanic anoxic events plasm. This could follow from the ingestion of food
documented by Bralower et al. (1994). An anoxic particles larger than those ingested by the ancestral
or dysoxic water column in the many parts of the Blefuscuiana and this would result (as suggested in
World Ocean would have prevented the earlier general terms by Banner and Desai, 1988) in the
dispersal of hedbergellids (and many also have ability of Hedbergella and Ticinella to thrive in
prevented their fossil preservation). waters deeper than those inhabited by the surface-
The Late Aptian was also a time of turnover in dwelling Blefuscuiana. The increase in depth and
the calcareous nannofossil record. Erba (1994) has probable decrease in the partial pressure of dis-
observed a marked turnover in nannoconid species solved oxygen could lead to an increase in the size
at the Early Aptian/Late Aptian boundary which of the test perforations, while the large size of par-
she terms the 'nannoconid crisis'. Large nanno- ticles demanded their disaggregation by muricae.
conids become extinct and are replaced by new, Certainly, the evolution of the macroperforate (and
deeper-water dwelling species. This may relate to muricate) test led immediately to the occupation of
the same events noted above which we cite as the oceans worldwide.
Plate 11.1

Figures 1-3 Hedbergella planispira (Tappan, 1940). Cenomanian, Grayson Fonnation, Danton County, Texas (from
Robaszynski and Caron, 1979, pI. 27, figs 3a--c), topotype donated by A.A.H. Wonders to the Museum National
d'Histoire Naturelle, Paris, no. FG 604: dorsal, peripheral and ventral views (x21O), showing macroperforations
and muricae all over the test.
Figure 4 Hedbergella trocoidea (Gandolfi, 1942). Late Aptian, Simferopo1, Crimea, deposited in the Geological Facu1ty of
Moscow State University, Russia: dorsal view (x155), showing macroperforations and muricae on the early whorls.
Figures 5-7 Praeglobotruncana delrioensis (plummer, 1931). Cenomanian, Grayson Fonnation, Texas (from Robaszynski and
Caron, 1979, pI. 43, figs 2a--c), topotype donated by J. Postuma to the Museum National d'Histoire Naturelle,
Paris, no. FG 640: peripheral, dorsal and ventral views (xl60), with macroperforations and muricae concentrated
and fused near the periphery.
Figures 8-10 Praeglobotruncana stephani (Gandolfi, 1942). Cenomanian, de la Breggia, Tessin, Switzerland (from Robaszynski
and Caron, 1979, pI. 48, figs 1a--c), topotype donated by M. Caron to the Museum National d'Histoire Naturelle,
Paris, no. FG 649: peripheral, dorsal and ventral views (xllO), with macroperforations and muricae concentrated
and fused near the periphery.
 12
The evolution of the
Planomalinidae
M.K. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

12.1 INTRODUCTION cameral sutures of the last whorl. These muricae,

on the periphery of the test, developed into the
Alanlordella evolved independently from unique, peripheral muricocarina of Planomalina
Blowiella in the Late Aptian (Figure 5.2) and itself, in the Late Albian.
rapidly gained macroperforations, as in A. bento- The early Alanlordella, A. bentonensis
nensis (Morrow, 1934). Just as in the evolution (Morrow) followed biostratigraphically by A.
of Hedbergella from Blefuscuiana (Chapter 11), banneri BouDagher-Fadel (1995), had about seven
the macroperforations evolved earlier than the to eight chambers in the last whorl and a normal,
muricae. The earliest Hedbergella of the Albian interiomarginal, symmetrical, single aperture
were often macroperforate but not yet muricate which extended from one umbilicus to the other.
(e.g. H. punctata, Michael, 1973; H. intermedia However, in the Late Aptian of the Caribbean and
Michael, 1973; H. modesta (Bolli) of Sigal, 1979; Central America, species evolved in which certain
H. globigerinelloides (Subbotina) of Sigal, 1979), individuals divided this aperture into two separate,
and some of these probably represent direct interiomarginal openings, each facing one of the
descent from microperforate, smooth immediate two umbilici. An obvious example was the so-
ancestors (e.g. the true 'Praeglobotruncana' called Biglobigerinella barri Bolli, Loeblich and
modesta Bolli, 1959, and 'Globigerina' globiger- Tappan (1957), first described from the Maridale
inelloides Subbotina, 1949) which are referable to Formation of Trinidad, in which many of the
Blefusciuiana. Similarly, A. bentonensis, which paratypes were found by its authors to become
first appeared in the Late Aptian but persisted until terminally biserial, just like the specimen which
the end of the Early Cenomanian (Figure 12.1), they selected as holotype (refigured here in Plate
became macroperforate but still lacked muricae; 12.2, Figs 4, 5). Such macroperforate, terminally
its immediate descendant, A. banneri, developed biserial species occurred many times in the Late
muricae on the early chambers of the last whorl, Cretaceous, and had been found in sedimentary
and evolved into A. praebuxtorji (Wonders) in rocks as young as Maastrichtian (Bolli, Loeblich
which the muricae were widespread in the inter- and Tappan, 1957, p. 25). We consider that it is
 -t.
-t. -t.
II) II) CD
-t. .... i:
.., m

APTIAN ALBIAN CENOMANIAN ,.

G)
m
EARLY LATE EARLY LATE EARLY LATE

Alanlordella apt/ens/s
A.banner/
A. bentonensls
A. praebuxtorfl
"B/glob/gerlnella" bam
Planomallna buxtorf/

- P. chen/ourens/s

Figure 12.1 Stratigraphical distribution of Early Cretaceous Alanlordella, 'Biglobigerinella', and Planomalina. Chronostratigraphy and Absolute Time Scale after
Gradstein et al. (1995).
 Systematics 211

probable that each Late Cretaceous 'species' of macroperforations and, soon afterwards, septal
Biglobigerinella evolved separately from different muricae. Then (probably only in the area of the
species of Planohedbergella gen.nov. (Appendix United States) in the latest Aptian and Albian, this
1), and, as they appear to be confined to the stock develops terminal biseriality which is char-
Americas and westernmost Tethys, they may have acteristic of Biglobigerinella. However, these
arisen under local ecological conditions, where terminally biserial forms occur as stratigraphically
both uniserial and biserial forms of each 'species' high as the Campanian-Maastrichtian as Biglobi-
occurred in the same sample. gerinella multispina Lalicker (1948); such forms
were unique among the Hedbergellidae in their
terminal biseriality and in their latest Cretaceous
12.2 SYSTEMATICS range and must be distinguished on the generic
level from all of the rest.
Superfamily GLOBIGERINACEA Carpenter, Alanlordella should be placed in the Plano-
Parker and Jones, 1862 malinidae because it was planispiral, compressed
laterally, macroperforate and never developed
Family PLANOMALINIDAE Bolli, Loeblich perforation cones (which only occurred in the
and Tappan, 1957 Praehedbergellidae), but it frequently developed
muricae. When muricae appeared, they first
Genus Alanlordella BouDagher-Fadel, 1995
occurred on the earliest part of the last whorl adja-
Type species Alanlordella banneri BouDagher- cent to the aperture, as in Alanlordella banneri,
Fadel, 1995 where, it is suggested, they could disintegrate the
food particles carried by the extrathalamous cyto-
Diagnosis: Test planispiral, nearly involute; plasm directly to the aperture. By the Cenomanian,
chambers globular, sutures depressed; wall muri- almost all species of these planispiral forms possess
cate in early whorls, macroperforate, the macro- muricae on the intercameral sutures of all chambers
perforations becoming sunk into perforation of the last whorl and a peripheral muricocarina
depressions and the rims of these depressions fus- (or muricate 'keel'). There is a gradual spread of
ing to give the later chambers a cancellate texture. the muricae during Aptian-Albian-Cenomanian
Periphery rounded, sometimes compressed in the evolution, from a partial muricocarina, as in
early stages but never developing a keel. Aperture Alanlordella, to the fully keeled (fully muricocari-
interiomarginal, equatorial, symmetrical, extending nate) Planomalina. The transition between these
laterally into each umbilicus; final aperture and genera is essentially gradual. They are retained
relict apertures are usually furnished with portici. because of their stratigraphical value. The bound-
ary between them is arbitrary and they have been
Remarks: The genus at present is considered to differentiated because of their utility in describing
have four species: A. banneri BouDagher-Fadel, the phylogenetic evolution of the Globigerinacea in
1995 (the type species), A. aptiensis (Longoria, this part of the Cretaceous.
1974), A. bentonensis (Morrow, 1934) and A. The Planomalinidae are planispiral and possess
praebuxtorji (Wonders, 1975). Alanlordella is strong muricae (at least on the early chambers of
different from Globigerinelloides in being macro- the last whorl) and can develop a peripheral
perforate and muricate and from Planomalina in muricocarina. Alanlordella lacks a muricocarina
lacking a muricocarina. It does not show the bi- except on the earliest chambers of the last whorl,
seriality of Biglobigerinella or the extensive while true Planomalina has a muricocarina on all
muricae, which develop over all of the test, of chambers of the last whorl.
Planohedbergella gen.nov. (Appendix 1). Alan-
lordella developed in the Late Aptian from the Range: Latest Aptian to Cenomanian (the true
planispiral, microperforate (sometimes with per- hedbergellid Biglobigerinella, unlike the plano-
foration cones) Globigerinelloides by acquiring malinid homeomorph 'B.' barri Bolli et al. (1957),
212 The evolution of the Planomalinidae

occurring through the Late Cretaceous). Alan- Alanlordella aptiensis (Longoria, 1974)
lordella appears again in the Taylor Formation of
(Plate 12.2, Figs 1-3; Figure 12.1)
Texas in the shape of Globigerinelloides bollii
Pessagrto (1967) which is more compressed than 1974 Globigerinelloides aptiense Longoria (sic),
A. bentonensis (Morrow) but otherwise very pp. 79, 80, pI. 4, figs 9, 10, pI. 8, figs 4-6,
similar (see McCarthy, in preparation); we do not 17, 18.
know the immediate origin of A. bollii as there are
no specimens yet known of Alanlordella between Remarks: This species has five to six inflated
the Cenomanian and Campanian. chambers in the last whorl, increasing gradually
AlanlordeUa banneri BouDagher-Fadel, 1995 in size, a strongly lobate periphery and wide
umbilici. Although it has no muricae, its wall is
(Plate 1.1, Fig. 6, Plate 12.1, Figs 1-9; Figure distinctly macroperforate and the species is clearly
12.1) an early form of Alanlordella. It is similar to A.
1987 Globigerinelloides caseyi (Bolli, Loeblich bentonensis (Morrow) but has much broader
and Tappan), Ben Haj Ali, pI. 4, fig. 12. umbilici, more deeply depressed intercameral
1987 Globigerinelloides sp., Ben Haj Ali, pI. 4, sutures, and fewer chambers in the last whorl (five
fig. 13. to six, compared to seven to eight).
1995 Alanlordella banneri BouDagher-Fadel,
p. 150, pI. 2, figs 10-12. Range: Within the Late Aptian. This species
was first described from the G. ferreolensis Zone
Diagnosis: Test large, about 460-550 fLm at its (Gargasian) of southeastern France.
maximum diameter, planispiral, early whorls
Alanlordella bentonensis (Morrow, 1934)
compressed; eight chambers in the last whorl,
slightly inflated, narrow, higher than broad, (Plate 12.3, Figs 1-9; Plate 12.4, Figs 1-9;
increasing gradually in size, sutures slightly Figure 12.1)
curved depressed; wall macroperforate with sharp
muricae on the early whorls, the macroperforations 1934 Anomalina bentonensis Morrow, p. 201 , pl.
becoming deep and the rims of these depressions 30, figs 4a, b.
fusing to give a cancellate texture; aperture interio- 1957 Planomalina caseyi Bolli, Loeblich and
marginal, equatorial, symmetrical, extending later- Tappan, p. 24, pI. 1, figs 4, 5.
ally into small umbilici, relict umbilical apertures 1964 Globigerinelloides caseyi (Bolli, Loeblich
furnished with thin portici. and Tappan), Low, pp. 122, 123.
1966 Planomalina (Globigerinelloides) caseyi
Remarks: This species differs from A. bento- (Bolli, Loeblich and Tappan), Salaj and
nensis (Morrow) in having a higher number of Samuel, p. 161, pI. 6, fig. la, b.
(narrower) chambers, a muricate test with very 1967 Globigerinelloides bentonensis (Morrow),
concentrated macroperforations. It differs from Pessagno, p. 275, pI. 76, figs 10, 11.
Alanlordella praebuxtorji (Wonders) in lacking 1967 Globigerinelloides caseyi (Bolli, Loeblich
muricocarina on the periphery of the early and Tappan), Pessagno, p. 276, pI. 49, figs
whorls, and in the much deeper and larger macro- 2-5.
perforations. 1975 Globigerinelloides caseyi (Bolli, Loeblich
and Tappan) emend., Wonders, p. 88, pI.
Range: Early to Late Albian. This species was 1, figs 3a, b, p. 89, text-fig. 4, figs la, b.
first recorded in the Early Albian of Tunisia 1977 Globigerinelloides bentonensis (Morrow),
(BouDagher-Fadel, 1995). Similar specimens had Masters, p. 404, pI. 10, figs 2, 3.
been figured by Ben Haj Ali (1987) (under 1977 Globigerinelloides cushmani (Tappan),
different names) from Tunisia, this time from the Masters, pI. 10, fig. 1.
Late Albian. 1977 Globigerinelloides bentonensis (Morrow),
 Systematics 213

Carter and Hart, pp. 27, 28, pI. 1, fig. 11; Loeblich and Tappan's original illustrations of
pI. 2, figs 19, 20. caseyi, and with those of 'G' bentonensis
1980 Planomalina (Globigerinelloides) bento- (Morrow). All are considered conspecific. More-
nensis (Morrow), Salaj, Figs 11, 13,21-23. over, the Tunisian and the English specimens are
1981 Globigerinelloides bentonensis (Morrow), identical in that the last formed chambers are still
Hart et al., p. 198, pI. 7.13, figs 7-9. microperforate (like the ancestral Blowiella) but
1987 Globigerinelloides bentonensis (Morrow), the early chambers of the last whorl have become
Ben Haj Ali, p. 86. macroperforate. This species is transitional from
1987 Globigerinelloides eaglefordensis (More- microperforate Blowiella to the macroperforate
man), Ben Haj Ali, pI. 4, figs 14, 15. Alanlordella. In the latter genus muricae begin on
1995 'Blowiella' bentonensis (Morrow), the early chambers of the last whorl and are absent
BouDagher-Fadel, p. 145, pI. 3, figs 1-9, on the later chambers. This evolution is clearly
pI. 4, fig. 6. protogenetic in that the morphological features of
the adult ancestors are recapitulated in the latest
Diagnosis: Test planispiral, about 210-360 I-Lm stages of life while the newly evolved characters
at its maximum diameter, biumbilicate; six to (macroperforations and muricae) appear in the
seven chambers in the last whorl, globular, almost early stages of ontogeny during evolution. They
as high as broad, increasing rapidly in size; sutures later extend to all growth stages, and with the
radial and depressed; aperture interiomarginal, acquisition of muricae become true Alanlordella.
equatorial, symmetrical, extending laterally into On the other hand, the umbilicus of A. bentonensis
each umbilicus for half of the last whorl, furnished is wider than in typical Blowiella but it has
with portici; wall smooth, not muricate, but not reached the great width of typical Globi-
macroperforate in the early part of the last whorl; gerinelloides.
the macroperforations are visible only in the early Carter and Hart (1977) and Hart et al. (1981)
part of the whorl, but the later chambers retain the were correct in considering that A. bentonensis
microperforations of this species' ancestors. arose from Blowiella, but the species of immediate
ancestry has yet to be determined.
Remarks: In rare specimens, later in the Early
Albian, the antepenultimate chamber develops a Range: With the Late Aptian-Early Ceno-
tubulospine (Plate 12.1, Fig 9) like Schackoina. manian. 'B.' caseyi was first described from the
Anomalina eaglefordensis Moreman (1927), Late Albian to Early Cenomanian strata (Carter
considered by Loeblich and Tappan (1961) to be and Hart, 1977 and Hart et al., 1981) but in Tunisia
a senior synonym of this species, was subse- it occurs in beds as old as Late Aptian. This is
quently confirmed as a benthonic species (Low, another example of the stratigraphically earlier
1964; Pessagno, 1967) and all the planktonic appearance of species in low latitudes, and a later
individuals referred to it were placed by them in appearance in more northerly areas. In Tunisia, it
Globigerinelloides caseyi (Bolli, Loeblich and was found in the Late Aptian, in the Early Albian,
Tappan), a species first described from the Albian and in the Albian-Early Cenomanian.
Gault Clay of England. However, after examina-
tion of topotype material, Carter and Hart (1977, Alanlordella praebuxtorfi (Wonders, 1975)
p. 28) rejected the name caseyi using G. bento- (Plate 12.5, Figs 1-9, Plate 12.6, Figs 1-6;
nensis for the Late Albian and Cenomanian forms Figure 12.1)
found in the British succession.
A suite of specimens from Tunisia and from 1975 Planomalina praebuxtorfi Wonders, pp.
the Gault Clay Formation, Arlesey brick pit, 90, 91, pI. 1, figs 1, 2, text-fig. 4, fig. 2.
Cambridgeshire (the type locality of G. caseyi) are
illustrated in Plate 12.3, Figs 1-9, Plate 12.4, Figs Remarks: This species is distinguished in hav-
1-9) and these have been matched with Bolli, ing depressed sutures, muricae concentrated near
214 The evolutiou of the Plauomaliuidae

the periphery and on the early sutures of the earlier by Bolli, Loeblich and Tappan (1957). It was later
part of the last whorl, the muricae fusing into a recorded by Longoria (1974) from Mexico and
peripheral muricocarina on the early chambers of France.
the last whorl, the later chambers being com-
pressed but not carinate. Genus Planomalina Loeblich and Tappan, 1946
This species is the intermediate stage between
Type species Planomalina apsidostroba Loeblich
A. bentonensis (Morrow) and Planomalina
and Tappan, 1946 = Planulina buxtorji. Gandolfi,
buxtorji. (Gandolfi).
1942

Rauge: Late Albian. Wonders' original material Praehedbergella evolves into the biumbilicate,
came from Jaen Province, southern Spain. planispiral Blowiella (Early Barremian-Late
Specimens also of Late Albian age from ODP sites Aptian), which in tum gives rise to the broadly
in the SE Indian Ocean have been kindly made clavate genus Claviblowiella (in the Early Aptian)
available to us by Dr A. Holbourn. and then to a younger, broadly biumbilicate form
with many more chambers (Globigerinelloides) in
Genus Biglobigerinella Lalicker, 1948 the beginning of the Late Aptian. This, in tum,
Type species Biglobigerinella multispina evolves into the planomalinid macroperforate
Lalicker, 1948 Alanlordella and then progressively in the latest
Aptian into the macroperforate Planomalina by
'Biglobigerinella' barri (Bolli, Loeblich and
acquisition of a peripheral keel (muricocarina).
Tappan, 1957) Planomalina ranges from the latest Aptian into
(Plate 12.2, Figs 4-9; Figure 12.1) the Cenomanian. This phylogeny is shown in
Figure 5.2.
1957 Biglobigerinella barri Bolli, Loeblich and
Tappan, p. 25, pI. 1, figs 13-18b. Planomalina buxtorfi (Gandolfi, 1942)
1974 Globigerinelloides barri (Bolli, Loeblich
and Tappan), Longoria, pp. 80, 81, pI. 4, (Plate 12.6, figs 7-9; Figure 12.1)
figs 1-3, 8, 14, pI. 5, figs 9-16, pI. 27, 1942 Planulina buxtorji. Gandolfi, p. 103, pI. 3,
fig. 19. figs 7a-c.
1946 Planomalina apsidostroba Loeblich and
Remarks: This species has eight to ten globular Tappan, p. 258, pI. 37, figs 22, 23.
chambers in the last whorl; often, but not always, 1975 Planomalina buxtorji. (Gandolfi), emend.
with biserial final chambers. The wall is macro- Wonders, pI. 1, fig. 4, text-fig. 4, figs 3a,
perforate and the surfaces of the earliest chambers b, 4a, b.
of the last whorl are muricate. 1985 Planomalina buxtorji. (Gandolfi), Caron,
Biglobigerinella with biserial final chambers p. 65, Figs 29.1-2.
occurred randomly in the Late Cretaceous;
however, it is not cosmopolitan. The Late Remarks: This species is distinguished by
Cretaceous species probably evolved from having eight chambers in the last whorl (increasing
planispiral, uniserial forms of Planohedbergella gradually in size as added), and by being fully
gen.nov. (Appendix 1); their ancestry is unknown, muricocarinate (possessing an entire peripheral
and they were probably unrelated to 'B'. barri keel round the margin and extending into the
(which we believed to have evolved from Aptian sutures).
Alanlordella), hence the informal use of the genus
in this case. Range: Late Albian to Late Cenomanian. The
type locality of P. apsidostroba (type species of
Range: Late Aptian. This species was first the genus but a synonym of P. buxtorji.) was in
described from the Maridale Formation of Trinidad the Main Street Limestone in Johnson County,
 Concluding comments 215

Texas, dated on ammonites as Albian- have retained this species in the fully muricocar-
Cenomanian boundary. Wonders' specimens were inate Planomalina, as we have said before
from the Late Albian-Early Cenomanian of (p. 211), the transition between these genera is
southern Spain. Gandolfi's material came from the essentially gradual and arbitrary. Nevertheless this
Late Albian of the Breggia River, Switzerland. phylogeny is at variance with that suggested by
Our figured specimens come from the Late Albian Caron (1985) who would derive P. cheniourensis
of the Lindi River, Tanzania (ex British Petroleum from Globigerinelloides algerianus Cushman
Microfossil Reference Collection). and Ten Dam by progressive acquisition of the
peripheral keel.
Planomalina cheniourensis (Sigal, 1952)
(Plate 12.6, Figs 10-13; Figure 12.1) Range: Latest Aptian. This species was first
described from the latest Aptian of northern
1952 Planulina cheniourensis Sigal, p. 21, text- Algeria.
fig. 17.
1985 Planomalina cheniourensis (Sigal), Caron,
p. 65, Figs 29.3-4. 12.3 CONCLUDING COMMENTS

Remarks: This species differs from P. buxtorji The planispiral taxa described in this chapter
(Gandolfi) by its more numerous chambers (twelve developed a phylogeny which culminated in the
chambers in the last whorl) and absence of periph- unique genus Planomalina, the only genus of the
eral muricocarina on the last three chambers. order Globigerinina ever to develop a peripheral
According to our phylogeny (Figure 5.2), this muricocarina (Figure 5.2). P. buxtorji is a very
species must have evolved from a macroperforate useful biostratigraphic index for the Late Albian,
Alanlordella (which, as we have seen, can have and it is very possible that this family did not
muricocarina on the earliest chambers of the last survive beyond the Cenomanian - especially as
whorl), but presumably not from A. bentonensis, there is now evidence that 'Biglobigerinella' barri
from which the P. buxtorji lineage is thought to (Plate 12.2, Figs 4-9) has a different ancestry from
have derived (see further discussion under A. prae- that of Biglobigerinella multispina Lalicker, the
buxtorji (Wonders), pp. 214, 246). Although we type species of this genus.
Plate 12.1

Figures 1-9 Alanlordella banneri BouDagher-Fadel, 1995. Early Albian, sample 88170, Beauvoir-ill Well, Tunisia, deposited
in The Natural History Museum, London, nos PF 53138, PF 62949 and PF 62950. 1,2, holotype (pF 53138),
opposite sides showing macroperforate wall surface and muricae on the early whorls (x85); 3, enlargement of wall
of holotype to show deep pores (xlO20), 4, 5, opposite sides of paratype (pF 62949) (x90); 6, enlargement of the
wall surface (x440). 7, 9, side and peripheral view of paratype (pF 62950) (x95); 8, enlargement of the wall
surface (x880).
Plate 12.2

Figures 1-3 Alan/ordella aptiensis (Longoria, 1974). Late Aptian (Gargasian), G.ferreolensis Zone, SE France (from Longoria,
1974, pI. 4, figs 9, 10; pI. 8, fig. 6), deposited in the US National Museum of Natural History, Washington, DC,
nos 185027, 185028: I, 2, holotype (185027), side view showing a wide umbilicus and a macroperforate test, and
peripheral view (x175); 3, paratype (185028), side view (x115).
Figures 4-9 'Biglobigerinella' barri (Bolli, Loeblich and Tappan, 1957). Aptian, Maridale Formation, Trinidad (from Bolli,
Loeblich and Tappan, 1957, pI. I, figs 14, 18a, b), deposited in the US National Museum of Natural History,
Washington, DC, nos P 4543, P 4544b: 4,5, holotype (P 4543), side and peripheral views showing biserial final
chambers (x95); 6, paratype (P 4544b), side view of specimen with uniserial final chamber (xl00). 7-9, Late
Aptian, La Peiia Formation, Mexico (from Longoria, 1974, pI. 5, figs 12-14), sample FL-373: 7,9, side views;
8, peripheral view (all xI25).
Plate 12.3

Figures 1-9 Alanlordella bentonensis (Morrow, 1934). Albian, Gault Clay, brick pit, Arlesey, S England, topotypes of
Planomalina caseyi Bolli, Loeblich and Tappan (1957), deposited in The Natural History Museum, London, nos
PF 62929-PF 62937. 1,2,4-9, side views showing a smooth, non-muricate wall but macroperforate in the early
part of the last whorl (1, x180; 2, xl40; 4, x180; 5, xl60; 6, x170; 7, x170; 8, x145; 9, x185); 3, peripheral view
showing equatorial aperture (xI80).
Plate 12A

Figures 1-9 Alanlordella bentonensis (Morrow, 1934). 1-3, Albian, Gault Clay, brick pit, Arlesey, S England, topotypes of
Planomalina caseyi Bolli, Loeblich and Tappan (1957), deposited in The Natural History Museum, London, nos
PF 62938, PF 62939: 1, side view (xI65), 2, enlargement of the surface showing a smooth, non-muricate wall
with regular perforations (x1l25); 3, side view (xI45). 4-8, latest Early Aptian, sample SS364, Jebel Chenanrafa,
Tunisia, deposited in The Natural History Museum, London, NHML PF 53l40-PF 53142: 4, 6-8, side views
showing portici visible in the umbilicus for about half of the last whorl (4,6, x205; 7, x175; 8, xl40), 5, periph-
eral view (x205). 9, Early Albian, sample SS538, Beauvoir-IV Well, Tunisia, deposited in The Natural History
Museum, London, no. PF 53144: side view (x270), showing the antepenultimate chamber with a tubulospine like
that of a Schackoina, showing the close relationship between the Blowiella-Alanlordella plexus and the
Schackoina-Leupoldina plexus.
Plate 12.5

Figures 1-9 Alanlordella praebuxtorfi (Wonders, 1975). 1-3, Late Albian, El Burrueco section, Jaen Province, S Spain (from
Wonders, 1975, pi. 1, figs la-c), deposited in the Utrecht Micropalaeontological Collections, State University of
Utrecht, The Netherlands, no. T 222: 1,3, side views of holotype showing the muricae fusing into a peripheral
muricocarina on the early chambers of the last whorl (xIl5); 2, peripheral view of same holotype, showing the
compressed test (x1l5). 4-9, Late Albian, south east Indian Ocean, presented by A. Holboum to The Natural
History Museum, London, nos PF 62951-PF 62956: 4,6,7, 9, side views of four specimens, showing muricae
concentrated near the periphery and the early sutures of the earlier part of the last whorl (4,6, x155; 7, xI20);
9, xl40). 5, 8, peripheral views of two further specimens, showing the equatorial aperture (5, xllO; 8, xl40).
Plate 12.6

Figures 1-6 Alanlordella praebuxtorji (Wonders, 1975). Late Albian, south-east Indian Ocean, presented by A. Holboum to
The Natural History Museum, London, nos PF 62957-PF 62962: 1-6, side views of six specimens, showing a
gradual increase of concentration of muricae near the periphery and the early sutures, forming an intermediate
stage between Alanlordella bentonensis (Morrow) and Planomalina buxtorji (Gandolfi) (1,2,4,6, x120; 3,5,
xI40).
Figures 7-9 Planomalina buxtorji (Gandolfi, 1942). Late Albian, Lindi area, Tanzania, deposited in the British Petroleum
Reference Collection at The Natural History Museum, London: 7, 8, side and peripheral views (xI05); 9, enlarge-
ment of the test to show macroperforations (x850).
Figures 10-13 Planomalina cheniourensis (Sigal, 1952). 10, II, Late Aptian, Jebel Oust, Tunisia (from Caron, in Bolli, Saunders
and Perch-Nielson (1985), Fig. 29; 3a, b): side view showing numerous chambers and the absence of periph-
eral keel in the last three chambers, and peripheral view showing a compressed test. 12, 13, Aptian, Oued
Cheniour section, N Algeria (from Caron, ibid., Fig. 29; 4a, b), holotype, deposited in the author's personal
collection, no. 101: side and peripheral views (x60).
 13
Aspects of the probable
physiology and ecology
of the Praehedbergellidae,
aided by studies of living
Globigerinacea
MK. BouDagher-Fadel, F.T. Banner
and J.E. Whittaker

13.1 INTRODUCTION as the living Globigerinacea, the Globigerinidae,

are worldwide in the oceans of today.
Can we learn anything about the ecology and A clear view of the palaeoecology of the
possible physiology of the Praehedbergellidae? Praehedbergellidae may be obtained from their
We can, by studying the distribution of praehed- fossil distribution and microfossil associations.
bergellid microfossil occurrences and, also but Their cosmopolitan distribution, in relatively deep-
more speculatively, by the nature of their living water, off-shelf sediments, immediately points to
analogues. Evolutionary convergence, to produce their holoplanktonic mode of life. It is possible
morphologically very similar tests, may imply that that the planktonic foraminifera began as mero-
similar habitats became occupied. We can explore plankton (Banner and Desai, 1988), with an adult
both of these possibilities. planktonic stage developed in order to enhance
With the sole exceptions of Globuligerina their reproductive strategies; the modem analogy
bathoniana, G. o;ifordiana and Favusella itself, would be with the tretomphaloid discorbids
the Favusellacea had a very restricted fossil distri- (Tappan and Loeblich, 1973; Banner, Pereira and
bution; both the earliest (Conoglobigerina) and the Desai, 1985; our Plate 2.1). It is also possible that
last (Ascoliella) genera of the Favusellacea had the planktonic foraminifera originated in response
very localized geographic distributions. to a warm climate, thermally stratified and,
In contrast, the earliest Globigerinacea, the perhaps, poorly oxygenated water (Tappan and
Praehedbergellidae, were widespread throughout Loeblich, 1973). The warm deep waters of Tethys
the Hauterivian-Aptian oceans and deep seas, just and the early Proto-Atlantic would have had lower
230 Physiology and ecology of the Praehedbergellidae

oxygen solubility than the cooler seas of today; are usually irregularly spaced. These genera have
only in the surface of the sea could oxygen achieve no spines or perforation pits; their surfaces are
high concentrations. This is discussed in more either smooth (Plate 13.1, Fig. 12) or are partly
detail below, but it is clear that such a distribution or wholly covered by low, irregular pustules (Plate
of oxygen would favour young organisms which 1.5, Figs 1,2; Plate 13.1, Figs 1--6, 10, 11). All
occupied the plankton. known extant microperforate genera are trocho-
spiral (although some fossil ones were planispiral).
In contrast, the other genera (in the families of
13.2 COMPARISON WITH LIVING the living Globigerinacea) have regularly spaced
MORPHOLOGICAL ANALOGUES perforations of about 10 ~m diameter and are
termed macroperforate (plate 1.5, Figs 4, 6). They
There is only one family of the living Globi- mayor may not bear spines or spinules and their
gerinacea (the Candeinidae) which contains small, surfaces may be encrusted or pitted; pits or spines
microperforate taxa - they are superficially pustu- may be present in the neanic growth stage, if not
late, not smooth (Plate 1.5, Figs 1,2, Plate 13.1, earlier or later. They may be trochospiral, plani-
Figs 1--6, 10, Tenuitellinata bradyi; Plate 13.1, spiral or streptospiral; these coiling modes are used
Fig. 11, Tinophodella glutinata) but their tests are to help identify the different families and genera.
otherwise very similar to those of the Praehed-
bergellidae. The subfamilies of the other extant
Key to the families of the
family, the Globigerinidae, contain tests which are
GLOBIGERINACEA
macroperforate and relatively large (Plate 1.5, Figs
3, 4, Globigerina bulloides; Plate 1.5, Figs 5, 6, 1 Tests microperforate, non-spinose, without
Orcadia riedeli). continuous encrustation, without perforation
The extant Globigerinacea have been reclassi- cones, but with blunt pustules
fied for this chapter, and it can be seen that the Candeinidae Cushman
biological characters of the subfamilies, as far as 2 Tests macroperforate, adults spinose, no perfo-
they are known, show remarkable distinctions ration cones; sometimes may be encrusted or
paralleling those indicated by skeletal differences. coated with smooth cortex
The classification of the Globigerinacea must be Globigerinidae Carpenter, Parker and Jones
based on what can be determined from the char-
acters of the test. The tests of most extant species Family CANDEINIDAE Cushman, 1927
have been illustrated and described by Saito et al.
Wall microperforate in adult, with perforations
(1981) and the genera by Banner and Whittaker
irregularly scattered; surface unencrusted, but
(in press). The biological characters of different
smooth or with scattered pustules, but no spines;
taxa have been summarized by Hemleben et al.
aperture (or apertures) interiomarginal, with
(1989). All this information is among that used in
narrow lips (or lips) of uniform breadth; test
this chapter. We recognize 22 genera of living
trochospiral. Never possesses the pointed muricae
Globigerinacea, which are listed below. About one
of the Favusellacea, or the perforation cones of
hundred species and subspecies are extant in
the Praehedbergellidae (Globigerinacea). We
modem seas.
recognize two subfamilies - the Globigerinitinae
and the Candeininae.
13.3 CLASSIFICATION OF LIVING
Subfamily GLOBIGERINITINAE Bermudez,
GLOBIGERINACEA
1961
Some genera (those of the family Candeinidae) Primary aperture single; dorsal sutural apertures
have tests which are microperforate: the perfora- absent or very few and sporadic. The species are
tions are about 1.5 ~m (or less) in diameter and characteristically highly eurythermal, thriving in
 Classification of living Globigerinacea 231

surface and subsurface uppermost waters from the the former may be broadest medially); coiling
tropics to subpolar seas. Some are believed to trochospiral or planispiral or streptospiral.
ingest fine vegetable debris, diatoms and chryso-
phytes. None is known to be carnivorous. Key to the subfamilies of the
GLOBIGERINIDAE
1 Test smooth except for distal, peripheral ends
Key to the genera of the
of chambers where thick spines, triradiate in
GLOBIGERINITINAE
section, are situated Hastigerininae Bolli,
1 Aperture intraumbilical 2 Loeblich and Tappan
I' Aperture intra-extraumbilical 4 I' Test smooth or punctate; spines, if present,
2 lacking bulla Tenuitellinata Li circular in section, and scattered over the
(Type species Globigerina whole test (at least in the juvenile) 2
angustiumbilicata Bolli) 2 Adult coated by cortical crust which narrows
2' With umbilical bulla 3 or blocks the exterior ends of the perforations
3 No accessory bullate tunnels Sphaeroidinellinae Banner and Blow
Globigerinita Bronnimann 2' Tests without a cortical crust (i.e., no
(Type species G. naparimaensis Bronniman cortex) 3
3' Infralaminal mUltiple apertures at end of 3 Tests trochospiral, sometimes becoming
bullate extension tunnels planispiral or pseudoplanispiral during
Tinophodella Loeblich and Tappan growth 4
(Type species T. ambitacrena 3' Test streptospiral 5
Loeblich and Tappan) 4 Surface of adult test spinose 6
4 Lacking bulla Tenuitella Fleisher 4' Surface of adult test non-spinose
(Type species Globorotalia gemma Jenkins) Globorotaliinae Cushman
4' With bulla Tenuitellita Li 6 Test large; surface cancellate
(Type species Globigerinita iota Parker) Globigerininae Carpenter, Parker and Jones
6' Test small; surface encrusted and smooth
Turborotalitinae Hofker
Subfamily CANDEININAE Cushman, 1927 5 Test smooth, adult wall without spines, aper-
ture single, interiomarginal
Early growth stage as in the Globigerinitinae, but
PuIleniatininae Cushman
adults with umbilicus closed and the single
5' Test punctate, adult thickly spinose, aper-
primary aperture replaced by many small, uniform
tures multiple, areal
apertures in each suture; surface smooth. Rare in
Orbulininae Schulze
tropical waters. Only one extant species, which
has never been studied biologically.
Candeina d'Orbigny Subfamily HASTIGERININAE Bolli, Loeblich
(Type species C. nitida d'Orbigny) and Tappan, 1957

Family GLOBIGERINIDAE Carpenter, Parker Key to the genera of the HASTIGERININAE

and Jones, 1862
1 Adult chambers radially elongate, clavate, often
Test macroperforate, with perforations regularly partly distally divided into two or more lobes;
and geometrically arranged (when not obscured by test initially planispiral but becoming strepto-
encrustation); surface may be pitted, with or spiral
without spines in adult, but may be smooth in adult Hastigerinopsis Saito and Thompson
(especially when encrusted) except for scattered (Type species H. digitiformans
pustules; apertures interiomarginal or areal (lips in Saito and Thompson)
232 Physiology and ecology of the Praehedbergellidae

I' Adult chambers not radially elongate 2 been studied biologically, but it is characterized
2 Coiling planispiral Hastigerina Thomson by possessing an additional, dorsal, sutural,
(Type species H. murrayi Thomson supplementary aperture. This genus is:
= Nonionina pelagica d'Orbigny) Sphaeroidinella Cushman
2' Coiling trochospiral (Type species Sphaeroidina bulloides
Orcadia Boltovskoy and Watanabe d'Orbigny excavata Parker and Jones)
(Type species Hastigerinella riedeli
Rogi and Bolli) Subfamily GLOBIGERININAE Carpenter,
Parker and Jones, 1862
Although Orcadia (Plate 1.5, Figs 5, 6) has not
yet been studied living, both Hastigerina and All species of the genera included here possess
Hastigerinopsis have been found to have a delicate calcitic spines radiating from the surfaces
calcareous test surrounded by a bubble-like of the adult test.
capsule of vacuolated cytoplasm, a feature not
known to occur in species of any other subfamily.
Key to the genera of the GLOBIGERININAE
Hastigerina has no algal, dinoflagellate or diatom
symbionts, although it may contain commensal 1 Adult test trochospiral 2
Pyrocystis robusta or P. noctiluca, but seems to I' Adult test planispiral 3
be exclusively carnivorous (the bubble capsule 2 With dorsal sutural supplementary apertures
may help to effectively digest captured prey); it Globigerinoides Cushman
also has a synodic lunar periodic reproduction (Type species Globigerina rubra
cycle (with disaggregation of the large, lobate d'Orbigny)
nucleus into micronuclei to form biflagellate 2' Without dorsal sutural supplementary
gametes), and massive calcitic spines (triradiate in apertures 4
section and barbed) along with cytoplasm flows to 4 Adult chambers subglobular
form rhizopodia which drag prey to the digestive Globigerina d'Orbigny
capsule. These features are unknown in other (Type species Globigerina bulloides
families, and may prove to characterize all the d'Orbigny
Hastigerininae. 4' Adult chambers radially elongate
All are of low latitudinal distribution, and Beella Banner and Blow
Hastigerinopsis, in particular, commonly inhabits (Type species Globigerina digitata Brady)
Deep or Intermediate Water layers. 3 Adult chambers subglobular
Globigerinella Cushman
Subfamily SPHEROIDINELLINAE Banner (Type species Globigerina aequilateralis
and Blow, 1959 Brady = G. siphonifera d'Orbigny
3' Adult chambers radially elongate
This subfamily was proposed to include genera
Bolliella Banner and Blow
which evolved during the Early Miocene and
(Type species Hastigerina (Bolliella)
which had adult tests coated in a thick, smooth
adamsi Banner and Blow
cortex of calcite; the cortex diminished or closed
the perforations and greatly thickened the aper- The trochospiral genera include both species
turallips. Tests which are broken show the surface which possess dinoflagellate photosymbionts and
of the younger whorls, below the cortex, to be those which do not; the former are restricted to
punctate and spinose. All the genera possessed the photic zone of the water column, while the
principal, primary apertures which were single, latter can live both in deeper water (down to 400
interio-marginal and intraumbilical in position and m). Even those which have symbionts may also
extent. be carnivorous (e.g. feeding on calanoid cope-
Only one, mono specific genus is extant, and this pods), but this is not universal. Globigerina occurs
is rare in subsurface, tropical waters; it has not in tropical to subtropical seas. Beella has never
 Classification of living Globigerinacea 233

been studied biologically, and seems to be con- Orbulina d'Orbigny

fined to the deeper waters of rather low latitudes. (Type species Orbulina universa d'Orbigny)
The planispiral genus Globigerinella possesses
Orbulina is distributed from tropical to temperate
two different photosymbionts, both of them
seas, but is much commoner in low latitudes (it
being chrysophytes. It occurs in tropical and
is not known to thrive at temperatures below
subtropical waters. Bolliella is much less common
12C). It is carnivorous, preying upon several
in these latitudes, and nothing is known of its
species of copepod and other organisms. However,
biology.
commensal dinoflagellates swim amidst its spines
and Gymnodinium is its mobile endophytosym-
Subfamily PULLENIATININAE Cushman,
biont (up to 23,000 such symbionts have been
1927
found inside one adult test).
The initial coils, like the adult ones, are strepto-
spiral; the chambers increasingly embrace Subfamily GLOBOROTALIINAE Cushman,
ventrally, so that the umbilicus becomes closed 1927
and the aperture becomes wholly extraumbilical.
In morphologically most advanced forms, the Key to the genera of the
chambers begin to embrace dorsally, so that GLOBOROTALllNAE
the last whorl covers much of the preceding
1 Equatorial periphery carinate
whorls. The surface of the neanic chambers is
Globorotalia Cushman
punctate, perforation-pitted and spinose, but the
(Type species Pulvinulina menardii
surface of the adult chambers is encrusted and
(d'Orbigny) var. tumida Brady)
smooth (except for tubercles around the aperture).
l' Equatorial periphery not carinate 2
There is only one extent genus:
2 Surface smooth; adult chambers often elongate
Pulleniatina Cushman
Clavatorella Blow
(Type species Pullenia sphaeroides
(Type species Hastigerinella bermudezi Bolli)
(d'Orbigny) var. obliquiloculata
2' Surface rough; adult chambers not
Parker and Jones)
elongate 3
Pulleniatina occurs in tropical seas, but little is 3 Principal aperture umbilical-extraumbilical
known of its biology. It appears to be solely throughout adult growth
herbivorous, and has been found to contain Paragloborotalia Cifelli
both diatoms and chrysophytes (it may be that all (Type species Globorotalia opima opima
have been ingested, but some may be living Bolli)
endosymbionts) . 3' Principal aperture becomes intraumbilical
in last few chambers
Subfamily ORBULININAE Schultze, 1854 Neogloboquadrina
Bandy, Ferichs and Vincent
The coiling is intensely streptospiral, with the last,
(Type species Globigerina dutertrei d'Orbigny)
globular chamber completely embracing the
ventral side (or all) of the juvenile test. The The peripherally carinate (keeled) species (of
interiomarginal principal aperture of the nepionic Globorotalia) develop smooth tests which may
and neanic stages is replaced by a multitude of possess scattered, small tubercles but which are
rounded pores in the area of the last chamber (and not cancellate or punctate - unlike the unkeeled
also in the last inercameral sutures if the nepionic genera (such as Paragloborotalia) , which never
test is not completely enclosed by the ephebic, have smooth adult tests (except when encrusted,
adult chamber). The surface of the adult test is and that is rare). Micropalaeontological studies
covered with perforation-pits and spines. There is have shown that species of Globorotalia repeat-
only one valid extant genus: edly evolved from species of Paragloborotalia;
234 Physiology and ecology of the Praehedbergellidae

Figure 13.1 A living species of Globorotalia (probably belonging to the G. cultrata (d'Orbigny, 1839) gr.), drawn by Mr JJ.
Wild from a surface specimen taken in a tow-net, off New Guinea, during the HMS Challenger Expedition (from Brady, 1884,
Fig. 17, p. 598). Although Brady supposed Wild's drawing to represent 'spines', no known Glororotalia has spines: (a) in these
positions; (b) so separated; (c) so thick; (d) so short; and (e) so curved. Rather, we believe them to be pseudopodia and this
remarkable figure is, to our knowledge, the first representation of the extrathalamous cytoplasm giving rise to pseudopodia on
the surface and near the periphery of a globorotaliid foraminifer.

the separate lineages have been given different their calcitic crusts readily distinguish them from
subgeneric names, but they are not employed here. juvenile tests of taxa of other families.
Paragloborotalia species are wider spread 1 Apertural lip weak or absent
latitudinally (being quite common in subpolar seas Berggrenia Parker
(e.g. P. incompta) to the tropics), but carinate (Type species Globanomalina praepumilio
Globorotalia is temperate to tropical only. Few Parker)
species have been studied biologically (Figure I' Apertural lip broad, making the final
13.1). Several species of Globorotalia ingest chamber ampullate, with small, accessory
diatoms or chrysophytes, but are not known to infralaminal apertures at margin of ampulla
possess photosymbionts; it is probable that they (which may cover the umbilicus)
inhabit waters deeper than those occupied by Turborotalita Blow and Banner
the unkeeled taxa. Neogloboquadrina also ingests (Type species Truncatulina humilis Brady)
phytoplankton, and has no known symbionts;
Species of Turborotalita carry endophotosym-
it is commonest in tropical waters but smaller
bionts (often chrysophytes) and live in the photic
forms (ecophenotypes?) range into temperate
zone for part, at least, of their lives; it is possible
waters.
that the adults live at greater depths and there
develop thicker calcitic crusts to increase the
Subfamily TURBOROT ALITINAE Hofker,
smoothness and heaviness of their tests. It has a
1976
transglobal, tropical-subtropical distribution.
The species of this subfamily are all small - the Berggrenia is known only from tropical latitudes
adults may be half the diameter of adults of other of the eastern Pacific, and it has not been studied
subfamilies - but their numbers of whorls and biologically.
 Palaeoecology 235

13.4 CONCLUSIONS FROM the survival of the organism. The test can have
COMPARISONS WITH LIVING had little or no defensive value, because predators
ANALOGUES who ingest plankton (like the modem whales and
many fish) would not have been deterred by such
Unfortunately, little is known of the biology of a small calcareous object.
the skeletal analogues of the Praehedbergellidae, However, the test was perforate, and these per-
which are the Globigerinitinae, a subfamily of forations have been shown, in many benthonic
the Candeinidae. Some Globigerinitinae are illus- rotalines, to act as links between the cytoplasm
trated on Plate 13.1; they are very similar to the which is inside each chamber (intrathalamous) and
praehedbergellids in their small size, thin walls that which forms a layer outside the test (extrathal-
and microperforations, but they never possess amous). It is from the extrathalamous cytoplasm
perforation cones though they often have surface that the pseudopodia arise, and they are 'granular';
pustules covering part or all of the test, and their the so-called granules are excreted particles being
apertural lips are narrow and symmetrical, never carried away from the test and captured organic
the asymmetrically long portici seen in the prae- particles being carried to the test for ingestion.
hedbergellids . When the captured particles reached the extrathal-
The Globigerinitinae are highly eurythermal, amous cytoplasm covering the test, they are
thriving in both warm and cool waters, but they dragged over the test surface to the umbilicus and
are confined to the surface and uppermost waters aperture for ingestion and digestion. The pseudopo-
of the world ocean. This latter character is clearly dia and the rest of the extrathalamous cytoplasm
similar to that already deduced for the Praehed- must also be the means with which the protozoan
bergellidae; the former may also be similarly gathers oxygen and excretes waste gases; these ions
comparable, but we do not know this yet. The pass through the perforations, even against con-
feeding habits of the Globigerinitinae must also centration gradients, into and out of the intrathala-
be comparable; the taxa are too small for them to mous cytoplasm (Angell, 1967; Sheehan and
be active, carnivorous predators, and some are Banner, 1972; Banner and Williams, 1973).
believed to ingest small diatoms and chrysophytes.
The Praehedbergellidae are likely to have had
The Early Cretaceous ocean and its plankton
similar feeding habits - the deeper water, exclu-
sively carnivorous habits of the larger, macroper- Caron and Homewood (1983) suggested that,
forate and spinose Hastigerininae and Orbulininae, while the open sea surface was colonized by 'prim-
for example, are totally alien to the Globigerini- itive' species during the Hauterivian-Aptian, more
tinae and also were likely to have been quite evolved, carinate species of Albian and younger
foreign to the Praehedbergellidae. times migrated into deeper oceanic waters. Hart
and Ball (1986) have related the evolutionary
appearance of Late Albian carinate genera
(Praeglobotruncana, rotaliporids) to the termina-
13.5 PALAEOECOLOGY
tion of Early Cretaceous oceanic anoxicity; they
suggest that anoxic subsurface waters had previ-
The function of the test
ously prevented deep water colonization by plank-
The calcareous skeleton of a planktonic fora- tonic foraminifera. We suggest that the
minifer must have had a function related to the microperforate tests of Jurassic and Cretaceous
separation of different parts of the cytoplasm of planktonic foraminifera were adequate to allow gas
the protozoan for different but related functions. and cation exchange between intrathalamous cyto-
Everything in the skeleton must have had a plasm and seawater in the well oxygenated sur-
purpose; the repetition of similar shapes and struc- faces of shelf seas and oceans, but the increased
tures, generation after generation, must have been test porosity which macroperforate evolution
because a skeleton of this kind was essential for would produce was necessary in order to maintain
236 Physiology and ecology of the Praehedbergellidae

efficient gas/cation exchange in less oxygenated either. Therefore, the relative amount of extrathal-
(and otherwise biogeochemically different) sub- amous cytoplasm in the Praehedbergellidae had to
surface waters. Also, in the thennohaline strati- be increased in other ways; one simple way of
fication of the Mesozoic ocean, vertical thennal doing this was to modify the chamber shape from
gradients, although weak, must have been created that of a sphere to something approaching a trape-
near-surface by solar heating and its turbulent dif- zoid, which has the maximum surface area for the
fusion to a relatively shallow Ekman depth; colo- minimum volume. Near-trapezoidal chambers have
nization of sub-surface cooler waters is likely to been known for decades to be present in
be enhanced by increased test porosity, as it is Schackoinidae with multiple tubulospines (e.g.
today in upper-water species from warm to cool Masella, 1960, plates 5, 6, 8) but a simpler and
latitudes (Frerichs et al., 1972). Therefore, the commoner way of reducing chamber-volume was
macroperforate HedbergeUidae and their descen- to elongate or flatten the chambers (or both).
dants are likely to have been more capable of pop- In such ways the relative proportions of extrathal-
ulating deeper, cooler and less oxygenated oceanic amous cytoplasm could be increased, and the
waters than were their microperforate ancestors. ability of the foraminifer to survive in increasingly
The absence of macroperforate Hedbergellidae dysoxic waters would be similarly enhanced. The
from the Late Aptian samples of areas such as broad, microperforate tubulospines of the Schack-
Speeton in England (compare the la Drome assem- oinidae would, themselves, increase the surface
blages described by Longoria, 1974) may be due area of the chambers, but the broader radial elon-
as much to the relative shallowness of the Late gation of the adult chambers of Lilliputianella,
Aptian seas in these areas as to their more northerly compared to the sub spherical chambers of
latitude, just as the porosity of Cenomanian Blefuscuiana, would also significantly increase the
Hedbergella and Rotalipora in Wyoming is less surface area of the chambers and therefore the
than in Texas (Frerichs and Ely, 1978). potential proportion of extrathalamous cytoplasm
and the ability of the foraminifer to survive in
waters lower in oxygen.
The significance of chamber shape and
oxycity
Oxygenation and chamber shape
If we consider solely the concentration of dissolved The surface water of the sea, stirred up by wind-
oxygen in the seawater, then, if the water is satu- generated waves, is nonnally saturated or super-
rated or supersaturated in oxygen, a relatively small saturated in oxygen by its physical solution from
volume of extrathalamous cytoplasm would be the air. It is in these most superficial layers of sea-
needed to maintain effective gas exchange. waters that we would find planktonic foraminifera
However, if the seawater becomes dysoxic, then with subglobular chambers. This is certainly the
relatively more extrathalamous cytoplasm, capable case with extant Globigerinacea. For example, in
of gas exchange, would be needed. A sphere has the Globigerininae, Globigerina with its subglob-
the greatest possible capacity for a minimum ular chambers lives near the sea surface, but
surface area; therefore, a spherical chamber would Beella, with its elongate chambers, inhabits deeper
have the greatest possible ratio of intrathalamous waters. In the Hastigerininae, Hastigerina itself,
to extrathalamous cytoplasm. This can be modified with subglobular chambers, floats at and very near
by producing vacuolated cytoplasm as in the the surface (John Murray went out in a rowing boat
Hastigerininae or extending the extrathalamous from HMS Challenger, in 1874, and collected
cytoplasm up thin spines as in the Hastigerininae Hastigerina from the sea surface with a spoon -
and Globigerininae; however, the Praehedber- as recorded by Wyville Thomson, 1877; Be and
gellidae and Favusellidae had no spines, and, by Anderson, 1976, used to capture individuals in jars
analogy with recent taxa as noted above, the while scuba swimming in the tropical west
Globigerinitinae (the closest analogue to the Atlantic!); in contrast, Hastigerinopsis, which has
Praehedbergellidae) had no vacuolated cytoplasm the most elongate chambers of any living
 Palaeoecology 237

Globigerinacean, is bathypelagic, being found at foraminifera which lived in the overlying waters.
depths greater than 1000 m (Be, 1965, p. 83). The depth-restricted living assemblages would be
These bathypelagic occurrences are far below the mixed together as dead tests on the sea floor.
depths of photosynthetic release of oxygen by Therefore, ignoring calcareous dissolution, the
planktonic algae, and are well into waters in which living assemblage (biocenosis) of the most super-
dysoxygenation is caused by metazoan respiration ficial water should always be represented in the
and decay. death assemblage (thanatocenosis) on the sea bed;
The Praehedbergellidae with subglobular cham- the biocenoses of slightly deeper, less oxygen-
bers (Gorbachikella, Praehedbergella, Blefus- saturated water, would be present as well if such
cuiana, etc.) are also likely to have inhabited both waters existed. Therefore, we would find assem-
the most superficial waters and also those which blages of the sub globular-chambered taxa in all
(although below the depths of sea/air mixing due normal marine samples, but these would be
to waves) diurnally gained dissolved oxygen from accompanied by species with elongate or flattened
algal photosynthesis. In contrast, the praehed- chambers in some samples only. The latter
bergellids with elongate chambers (Lilliputianella, species, with elongate or flattened chambers,
Claviblowiella, Leupoldina, etc.) are likely to have would not occur in samples which lack all taxa
occupied both the lower, fully oxygenated waters with subglobular chambers. This is, in fact,
as well as deeper waters with less oxygen, below what happens. Many areas yield samples with
the level of wave-air mixing and in which photo- the sub globular chambers only. For example, in
synthetic oxygenation also became much reduced. Tunisia, many Hauterivian-Early Aptian samples
have yielded Gorbachikella spp. but no other
Possible palaeobathymetry, estimated from planktonic foraminifera; these species have the
probable illumination most sub globular chambers of any Praehedber-
In the clear shelf-seas of the Early Cretaceous gellidae. In the central North Sea area, the
central Tethys (now in the off-shore of the United Barremian-Aptian succession contains (often
Arab Emirates), clean carbonate rocks were almost abundant) Praehedbergella and Blefuscuiana but
continuously deposited. These have now been only very rare Lilliputianella (Banner et aI., 1993).
sampled in hundreds of exploration wells. In a There are no known cases of Lilliputianella,
study using the principles of physical oceanog- Schackoina, Leupoldina or Claviblowiella occur-
raphy, as well as the fossil foraminifera and algae, ring in microfossil assemblages without the
to determine the levels of solar illumination in the subglobular-chambered taxa. The chamber-shape
Early Cretaceous seas, the palaeobathymetric of the Praehedbergellidae may be taken to char-
ranges of the foraminifera were estimated. It was acterize different levels in the upper waters of the
found that significant numbers of Praehedber- Early Cretaceous ocean.
gellidae first appeared at about 50 m below the An outstanding exception is the case of
Early Cretaceous sea surface, and abundance of Wondersella, which occurs in thin beds at the very
praehedbergellids did not occur until a depth of top of the Late Aptian Shuaiba Formation of off-
85 m or so was reached, at which depth fossil shore United Arab Emirates. This genus has the
calcareous algae were believed to cease significant most flattened and extended adult chambers of any
existence (Banner and Simmons, 1994). It is prob- praehedbergellid, but it occurs commonly and yet
able that in areas in which similar limestones were uniquely in samples from these beds (Banner and
developed in the early Cretaceous, the Praehed- Strank, 1987,1988). However, these sediments are
bergellidae occupied similar palaeodepths. clearly those of an anoxic-dysoxic sedimentary
basin. The micritic sediment has microlayeriug
Stratified biocenoses and the death but no signs of bioturbation, and contains no
assemblage benthonic microfossils - the bottom waters were
In the fossil assemblage, there should be an accu- anoxic. It is very probable that the upper waters
mulation of all the species of planktonic of this basin were markedly dysoxic: this is shown
238 Physiology and ecology of the Praehedbergellidae

by the fact that the waters were intolerable to all thalamous cytoplasm, as it deposits a new surface
other species of planktonic foraminifera and also lamella of calcite over the earlier test at each new
by the characteristic flattening and radial extension chamber formation. The penultimate chamber has
of the adult chambers of its only planktonic one extra lamella of calcite but earlier chambers
inhabitant - Wondersella. have more and more lamellae, thickening the test
and strengthening the cones. Automatically, this
Other ecological variables forms the strongest, most prominent cones nearest
Of course, this discussion so far has concerned to the aperture. The only analogues to perforation
only the effects of oxygen-saturation in the wave- cones in the Globigerinina are in some genera of
swept uppermost waters, the less-saturated waters the Heterohelicacea (Cassigerinella, Guembelitria,
below the wave-troughs, and the strongly dysoxic Pseudotextuiaria) but there are none in any
waters of very deep waters and those of environ- Holocene genera.
mentally exceptional basins. It does not consider In some benthic foraminifera, 'tubercles' (which
the effects on the globigerine test of the other, look the same as the perforation-cones of the
more readily observed variables such as salinity, Praehedbergellidae or the muricae of the
temperature, water density, illumination, food Hedbergellidae and their allies) are also concen-
supply, etc., but these must also have had marked trated on the test surface in the vicinity of the
effect on test shape and structure (as they do in aperture(s) and the umbilicus. Here, their function
modem planktonic foraminifera; Hemleben et al., seems to be the disaggregation by rasping of food
1989). Studies of living Globigerinacea suggest particles prior to their ingestion through the aper-
that oceanic variations in salinity do not limit their tures (Alexander and Banner, 1984) or digestion
biogeographic distribution (Bijma et al., 1990). in the umbilical cytoplasmic 'reservoir'
The same was probably true for the Praehed- (Alexander, 1985 MS). Is it not possible that the
bergellidae. With the exception of illumination, perforation cones and muricae of the apertural
the other variables are difficult to determine; areas of Globigerinacea, being similar in size,
temperature may be estimated from the radio- shape, density and distribution, had a similar
isotope ratios (e.g. oxygen isotope analysis; function?
Shackleton, 1967; Shackleton et al., 1985 and It is probable that some populations of particular
many other papers), but these have not been calcu- species of Praehedbergella and Blefuscuiana found
lated for the Early Cretaceous taxa. Differences in it more necessary to disaggregate their captured
ambient temperature may change test porosity (i.e. food particles than did other populations. This may
the number of perforations in any unit area of have been because the common food particulates
the test surface) but do not cause changes in the were larger in the waters where perforation cones
diameter of the perforations (Bijma et ai., 1990, were developed; such a case would arise either
especially plate 1). The same would have been because the organic detritus in the water had a
true for the Praehedbergellidae and Favusellacea. different biological origin, or because the waters
were slightly deeper, or both. In any case, region-
ally different populations of particular species of
The sculpture and structure of the test
Praehedbergellidae, possessing otherwise identical
surface
tests, are characterized by the presence or absence
The perforation cones of perforation cones, which must reflect the charac-
Species of the trochospiral Praehedbergellidae ters of the water masses in which these populations
either possess perforation cones on the surface of lived. Because we believe that the perforation cones
their tests, or they do not. When the cones are characterize particular environments, we name the
present, they become heavier and thicker towards bearers of the cones as distinctive subspecies which
the aperture; they become much lighter on the later inhabited those different environments, often in
chambers and may be absent on the last. This has different geographical areas. An example is Blefus-
been achieved by their formation by the extra- cuiana infracretacea s.s. and B. infracretacea
 Palaeoecology 239

occidentalis, the latter lacking the perforation cones pseudopodial extensions into the surrounding
so prominent in the former (Chapter 8). seawater. The pseudopodia would either have
extended irregularly or would have occupied a
The appearance of muricae and volume approximating to that of a sphere. If the
muricocarinae same volume is compressed into a disc, its surface
The large perforations are acquired first as the taxa area for prey capture or collection must be very
move into deeper water, then the muricae appear greatly increased. Therefore, species which could
gradually first near the aperture, then near the extrude the pseudopodia into such a disc must
periphery and sutures (in the evolution from have had considerable advantage in the collection
Blefuscuiana to Hedbergella, Globigerinelloides of available suspended particulate nutrients. This
to Alanlordella). As we have discussed, they could be the reason for the repeated success of
developed in order to dis aggregate the larger par- taxa which evolved muricocarinae (or other keels),
ticles which are found in these greater depths of and for the success of the Hedbergellidae and its
the water. descendants in supplanting the Praehedbergellidae
The function of muricae was almost certainly and Schackoinidae from their dominant position
the same as that discussed for perforation cones. in the biota.
However, the muricae occur with macroperfora- The Globotruncanidae, which descended
tions and we believe both to have been developed directly from Hedbergella, possessed tests which
in layers of seawater deeper than those inhabited were more elaborately sculptured. Globotruncana
by the Praehedbergellidae. The muricae charac- itself possessed two muricocarinae, one of which
terized the Planomalinidae and the Hedbergellidae extended into the cameral sutures of the dorsal
and their descendants. Unlike the perforation side and a ventral peripheral muricocarina which
cones, the muricae (which were formed between curved around the apertural face of each chamber
the perforations) could fuse together, to develop to the umbilical shoulder of each chamber of the
into murico-carinae. last whorl. Between these two muricocarinae there
With the acquisition of muricae in the Hedber- was an imperforate peripheral band which
gellidae and Planomalinidae, muricocarinae could extended into each imperforate septum. All the
develop (no keels are known in the Praehed- Globotruncanidae are extinct, so it is impossible
bergellidae and Schackoinidae). It is known that to observe the function of these structures, but we
the spines of living Globigerinidae are covered by must accept the principle that every structure had
extrathalamous extensions of the cytoplasm, capa- a function. There was no 'ornament' for these
ble not only of carrying vacuoles and transporting aesthetically limited organisms. We can surmise,
symbionts (Hemleben and Spindler, 1983, plate 1, from what has been noted above in other organ-
figs 3, 4) but also performing the feeding and isms, that the double muricocarinae of Globo-
exchange functions of exopodia when the latter are truncana was also associated with nutrition. If fans
lost (Adshead, 1980). The carinae of some preda- of pseudopodia radiated from the extrathalamous
tory, biconvex benthic Rotaliina (e.g. cytoplasmic layer, which covered the test, and
Amphistegina) have been observed to form the arose from each muricocarina, then they would
foundation of the 'take-off' of food-gathering drag organic particles to the muricacarinae where
pseudopodia (Banner, 1978). Is it not possible that they would become disaggregated. These disag-
the peripheral muricae, and, later the muricocari- gregated particles would then be channelled along
nae, of the Hedbergellidae and its descendants the imperforate band to the terminal face and then
functioned similarly, as a peripheral rim from to the umbilicus. This would mean that no food
which food gathering pseudopodia could extend in particles would impede the function of the perfor-
a disc-like fan? ations on the dorsal and ventral faces of the cham-
The smooth, nonmuricate, noncarinate Praehed- bers, but would ensure that simple cytoplasmic
bergellidae and Schackoinidae could have had no flow would direct the nutrient particles straight to
skeletal control on the distribution of sites for the umbilical aperture for their ingestion. Such
240 Physiology and ecology of the Praehedbergellidae

elaborate but efficient nutritional mechanisms rise and fall of the world sea-level during (at least)
were inevitably absent in the Praehedbergellidae the Mesozoic and Cenozoic (Haq, Hardenbol and
and the Schackoinidae, which is why those fami- Vail, 1987, 1988). It might be suggested that the
lies became dominated in the oceans by the resulting eustatic curves could indicate reasons
Hedbergellidae and their descendants. for the bursts of evolution which the Praehed-
The muricae of the Favusellacea and the Globi- bergellidae underwent during the Hauterivian-
gerinacea are morphologically indistinguishable Barremian and Late Aptian, and which the
but seem to be independent evolutionary devel- Hedbergellidae experienced in the Late Aptian-
opments. All the Favusellacea are essentially Albian (Hart, 1980). The eustatic curves (Haq et
muricate, from the Bajocian to the Cenomanian, al., 1988) suggest a global deepening of the seas
and the muricae (or the ridges derived from them) in the Barremian and in the Late Albian-
are distributed almost uniformly over each cham- Cenomanian, with eustatic shallowing in the
ber surface. In the Globigerinacea, muricae Valanginian and in the Aptain. These events do
appeared with the Hedbergellidae and the Plano- not relate convincingly to the evolutionary devel-
malinidae only from mid-Aptian times; then and opment of the Praehedbergellidae and its descen-
thereafter the muricae are concentrated in the dants, and this is not surprising. The eustatic
approaches to the apertures and on the test changes, with their coastal onlaps and offiaps, are
periphery. There is such a phylogenetic gap geologically seen in the neritic sedimentary
between the favusellacean and globigerinacean deposits of shelf seas; the planktonic foraminifera
developments that the muricae of each superfamily were the inhabitants of the upper and surface
cannot be homologous and may not even have waters of deep oceans. What happened in the
been analogous in function. sedimentary events of the continental shelves
could have been of little importance to them. If,
Umbilical cover plates and bullae in deep ocean, water depths changed by 250 m or
In some benthic Rotaliina (Haynesina, Rosalina) so when the oceanic depths were 1000 m or more,
partial closure by skeletal material of the umbili- the Praehedbergellidae of the uppermost seawaters
cal digestive cytoplasm (Alexander, 1985 MS) would have been unaffected. The evolution of the
seems to be advantageous for extrathalamous Praehedbergellidae would have proceeded inde-
digestion of disaggregated particles. Advanced pendently.
Hedbergellidae evolved analogous structures par- A similar conclusion must be reached regarding
tially to enclose their umbilici (e.g. Ticinella, the possible effects of the Mesozoic periods of
Rotalipora); this could be a similar partial enclo- anoxicity which have been recognized in sedi-
sure of an umbilical digestive 'reservoir' of cyto- ments bordering parts of Tethys and the proto-
plasm for similarly dis aggregated particles. Even Atlantic (Jenkyns, 1980; Schlanger and Jenkyns,
predatory, carnivorous globigerinids partially 1976; Kennett, 1982). Whatever may have
digest disaggregated prey in extrathalamous happened to the thermal deep-water circulation of
digestion vacuoles (Spindler et al., 1984). It is sig- the proto-Atlantic during its growth, the develop-
nificant that the Praehedbergellidae and Schacko- ment of anoxic periods there in temporarily
inidae, which never developed muricae, also never stagnant oceanic basins could have been events
developed extended portici to partly enclose the recognizable only in the sediments of the proto-
umbilicus. Atlantic. The narrow proto-Atlantic must have
been part of one world ocean which also contained
the broader proto-Pacific and proto-Indian oceans;
Praehedbergellid evolution in relation to
there is no reason to suppose that the thermal
sea-level change and anoxic periods
abyssal circulation of these latter oceans ever
Sequence stratigraphy and the development of experienced anoxic periods. The holoplanktonic
depositional models have suggested the cyclical Praehedbergellidae must have inhabited all the
 Palaeoecology 241

world oceans, and although the inhabitants of necessary increase in test porosity, but it also
the proto-Atlantic may have been affected by became advantageous to gather and disaggregate
anoxic periods, those of the rest of the world the larger, heavier particulates which had sunk
oceans would not have been. Therefore, the evolu- from the upper waters. Probably in order to
tion of the Praehedbergellidae, on a global scale, achieve this, aperturally-associated perforation
would not have been affected by local, even if cones (and in the Hedbergellidae and Planomalin-
regional, anoxicities. idae, muricae and, ultimately, modified umbilici)
were evolved.
Conclusion If more needs to be learned about the probable
palaeobiology of the Praehedbergellidae, then
We suggest that the Praehedbergellidae and more effort should be made in the biological
Schackoinidae lived at or near the sea surface study of its living analogues, the Globigerinitinae.
(where the warmth and oxygenation of surface Comparisons could then be suggested which
waters permitted survival with a low test porosity), could radically improve our understanding of
where the finest particulate nutrients remained the nature, life-styles, habits - and maybe, even,
in suspension. With the migration of Globigerin- evolution - of the Hauterivian-Barremian-Aptian
acea to deeper waters, not only was there a Globigerinacea.
Plate 13.1
Living members of the G10bigerinacea (Candeinidae; Globigerinitinae) are (often small) microperforate taxa (superficially pustu-
late, not smooth) with tests very similar to those of the Early Cretaceous Praehedbergellidae. All specimens from the collec-
tions of The Natural History Museum, London.

Figures 1-6 Tenuitellinata bradyi (Wiesner, 1931) (= Pylodexia uvula Ehrenberg nomen oblitum) a microperforate species with
scattered pustules concentrated near the intraumbilical aperture: 1, side view (x400); 2, 3, apertural and dorsal
view of same specimen (x500); 4, 5, two dorsal views and apertural view of specimen with a twisted (but still
intraumbilical) aperture (xI60).
Figures 7-9 Tenuitella anfracta (Parker, 1967): dorsal, peripheral and ventral views (x280).
Figure 10 Enlargement of surface of Tenuitellinata bradyi (Wiesner, 1931) to show microperforations (x2000).
Figure 11 Enlargement of surface of Tinophodella glutinata (Egger, 1893) to show pustules of a microperforate species
(xI250).
Figure 12 Enlargement of surface of Candeina nitida d'Orbigny, 1839 to show irregular microperforations (xI500).
 Appendix 1
Planohedbergella gen.nov.
M.K. BouDagher-Fadel, F.T. Banner,
J.E. Whittaker and Ll.D. McCarthy

last formed chambers of Planohedbergella, but are

Within the Hedbergellidae in the Late Cretaceous present only on the early chambers of the last
arose the new genus Planohedbergella. It is not whorl of Alanlordella, especially near the
unlike Alanlordella morphologically but is muri- periphery and in the early intercameral sutures, if
cate throughout. The evolutionary histories and they are present at all.
stratigraphic ranges of the two are also rather
different (Chapter 12). Remarks: Planomalina ehrenbergi Barr (1961,
p. 563, pI. 69, figs la, b; refigured here by SEM
Family HEDBERGELLIDAE Loeblich and on Plate App. 1, figs 1-3) has been chosen as the
Tappan, 1961 type-species for our new genus Planohedbergella
because this species has a weakly compressed test
Planohedbergella gen.nov. and about eight chambers in the final whorl - as
(plate Ap. 1, Figs 1-6) close as possible to the morphology of Alan-
lordella, so that comparisons between the two
Type species Planomalina ehrenbergi Barr, 1961 genera could be made fairly. The first publication
which dealt with the form we are calling
Diagnosis: Test calcitic, macroperforate, plano- Planohedbergella was that by Ehrenberg (1854)
spiral in adult; surface wholly muricate; aperture who named Phanerostomum asperum from the
umbilical-peripheral, interio-marginal, extending Late Cretaceous of Rugen, Germany. Topotypes,
between each umbilicus, over the periphery of from Ehrenberg's own sample, were refigured by
the penultimate whorl; umbilici are wholly or Masters (1980) who called them Globiger-
partially involute, and the aperture possesses a inelloides asper (Reuss).
porticus which is longest in the umbilici. [Gender:
feminine]. Comparative phylogenies: In the Plano-
malinidae, Alanlordella arose in the latest Aptian
Derivation of name: Refers to our phylogeny in from the praehedbergellid Globigerinelloides by
which it evolves from Hedbergella by becoming gaining macroperforations (A. bentonensis
planispiral. (Morrow, and then (in the Albian) by develop-
ing muricae also, first on the surface of the early
Differentiation: Differs from Alanlordella in chambers of the last whorl (A. banneri BouDagher-
being muricate throughout; muricae cover even the Fadel), and then in the intercameral sutures also
246 Appendix 1 Planohedbergella gen.nov.

(A. praebuxtorfi (Wonders)). The latter fuse to pro- done. Unlike the Alanlordella-Planomalina
duce a muricocarina on the periphery of the phylogeny, the evolving species of Plano-
adult test, which extends into the sutures, in hedbergella did not become more compressed and
Planomalina (P. buxtorfi (Gandolfi)). Throughout more multichambered; instead, the only descen-
this evolution, the evolving tests become more dant of Planohedbergella was the morphologically
compressed and acquire more and more chambers conspicuous but rare Biglobigerinella, in which
in the last whorl. the test became less compressed with growth, the
In contrast, in the Hedbergellidae, Plano- final chambers became very thick, the interio-
hedbergella arose in the Cenomanian from marginal aperture became divided by its closure
Hedbergella (Plate 11.1, Figs 1-4) by becoming over the periphery of the penultimate whorl (so
planispiral; this repeated evolution has been well that separate apertures opened in each umbilicus),
illustrated by Hofker (1960, plate 2). The earliest and then even the last chambers themselves
whorls of Planohedbergella may still possess low divided, to produce a brief, planospirally coiled,
trochospirality, but the adult final two or more biserial stage. Nothing at all like this ever hap-
whorls became clearly planispiral. Like the pened during the Alanlordella-Planomalina
ancestral Hedbergella, the chamber surfaces of phylogeny.
Planohedbergella were not only macroperforate In summary, although the morphological
but were uniformly and completely muricate; how- distinctions between the genera, Alanlordella and
ever, Planohedbergella never acquired a murico- Planohedbergella, may seem, at first sight, to have
carina either in the intercameral sutures or on its been small, their ancestries and evolutionary
periphery. The ancestral, trochospiral Hedbergella potential were totally different, as were their strati-
was able to fuse its perpipheral muricae to evolve graphic ranges. Needless to say, both genera
the muricocarinate Praeglobotruncana (plate 11.1, differed both morphologically and stratigraphi-
Figs 5-10), but the planispiral Planohedbergella cally from the praehedbergellid genus Globigerin-
never did, even though its had all the time of elloides, with which both the hedbergellid genera
the Late Cretaceous to do so, if it could have have frequently been confused.
Plate Appendix 1

Figures 1-6 Planohedbergella ehrenbergi (Barr, 1961). 1-3, 6, Coniacian/Santonian, Micraster coranguinum Zone, sample C42,
Culver Cliff, Isle of Wight, S England. SEM of the holotype figured by Barr (1961, pI. 69, figs la, b), deposited
in The Natural History Museum, London, no. P 44608: 1-3, opposite sides and peripheral view (><230); 6, enlarge-
ment of the wall surface showing macroperforations and muricae spread evenly (x800). 4, 5, Coniacian of S
England, from the British Petroleum Reference Collection in The Natural History Museum, London: opposite sides
(x31O).
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 Systematic Index

Suprageneric categories are printed in CAPITAL LETTERS, and genera and species in italics. The
principal reference for each species described in the book is marked with an asterisk (*). Page numbers
appearing in italics refer to figures and page numbers in bold refer to plates.

Alanlordella 56, 83,182,183,209,210,211-14,222, 'Biglobigerinella' 210,214

239,245,246 barri 210, 211, 214*, 215, 218, 219
aptiensis 210, 211, 212*, 218, 219 Biticinella ferreolensis 183
banneri 6, 7, 56, 209, 210, 211, 212*, 216, 217, Blefuscuiana 55, 56, 82, 83, 103, 105, 109, 110, 121,
245 122,123,124,132,133,161,166,203-5,209,
bentonensis 6, 7, 209, 210, 211, 212*, 213-15, 236-9
220,221,222,223,226 albiana 122, 123, 124*, 135,136,137
praebuxtorfi 209, 210, 211, 212, 213*, 225, 226, aptiana s.s. 6, 7, 122, 124, 125*, 126, 128, 133,
227,246 135,138,139
Amphistegina 239 aptiana imp lana 125
Anomalina aptiana orientalis 6, 7, 122, 124, 125*, 126, 133,
bentonensis 212 135,138,139,140,141
eaglefordensis 213 aptica 126, 129
lomeiana var. trocoidea 203 archeocompressa 103
Archaeokassabella biapertura 53, 54, 85, 92, 93 cf. aptica 128, 148, 149
Ascoliella 30, 55, 58,59,61-3,229 convexa 122, 123,126*, 142, 143
nitida 58, 61*, 62, 72, 73, 76, 77 daminiae 122,123,126*,144,145,158,159
quadrata 58,62*,72,73 excelsa s.s. 122, 123, 126*, 144, 145
scitula 58, 62*, 74, 75 excelsa cumulus 122,123,127*,133,144,145
scotiensis 55, 61, 72, 73 gorbachikae 122,123,127*,128,133,146,147,
voloshinae 58,62*,63,76,77 166
Asterohedbergella 166 hexacamerata 122, 124, 127*, 146, 147
asterospinosa 166 hispaniae 122, 123, 128*,135, 146, 147
infracretacea s.s. 80, 121,122, 123, 128*, 129,
Beella 232, 236 133,135,148,149,238
Berggrenia 234 infracretacea aptica 129
Biglobigerinella 211, 214, 216 infracretacea occidentalis 80, 122, 124, 128, 129*,
barri 209, 214 133,134,135,148,149,238
multispina 211, 214, 215 kuznetsovae 55, 122, 124*, 136, 137
260 SYSTEMATIC INDEX

laculata s.s. 103,122,123,129*,150,151, 152, globulifera 164

153 kuhryi 165
laculata alobata 122, 123, 129*, 150, 151 primare 131
mitra 122,123, 129*, 130, 150, 151 sigali 108
moulladei 123 tuschepsensis 105, 132
multicamerata 122, 123, 124, 130*, 133, 150, 151 Compactogerina 20, 21, 29, 30, 59
occulta s.s. 122, 124, 130*, 150, 151 stellapolaris 21, 29*, 44, 45, 46, 47
occulta perforocculta 122,124,139*, 150, 151 Conoglobigerina 4, 17-20, 21, 22, 24-6, 30, 53, 54,
occulta quinquecamerata 124, 125 57,62,63,85,229
praesimilis 103, 123, 129, 152, 153 araksi 22
praetrocoidea 122, 124, 130*, 131, 133,134, 152, avarica 4,17,21,22*,38,39
153,203,204 avariformis 17,21,23*,44,45,46,47
primare 122, 123, 131*, 154, 155 balakhmatovae 17, 18,21,22,23*,38,39
rudis s.s. 122, 123, 131*, 154, 155 bathoniana 27
rudis pararudis 122, 123, 131*, 154, 155, 156, 157 caucasica 19,21,24*,25,30,42,43,57
sp. 122, 123, 132*, 158, 159 conica 18, 21, 24*, 26
speetonensis s.s. 79, 122, 124, 131*, 132, 133, dagestanica 4,17,18,20*,21,22-6,36,37,38,
156,157 39
speetonensis tunisiensis 122, 124, 132*, 158, 159 gulekhensis 19, 21, 24*, 25, 30, 42, 43, 57
whittakeri 123, 126, 158, 159 jurassica 21, 25*, 44, 45, 46, 47
Blowiella 55, 82, 83, 110, 177, 178, 179, 182, 183, meganomica 4, 21, 25*, 40, 41
186,209,213,222 terquemi 18,21,24,26*
blowi S.s. 178, 179*, 183, 188, 189 CONOGLOBIGERINIDAE 20,21,29,30,59
duboisi 178, 179*, 188, 189 Cymbaloporetta
gottisi 178, 180*, 188, 189 cifellii 32, 33
maridalensis 178, 180*, 188, 189 sp. 32, 33
moulladei 178, 179, 180*, 190, 191 Cymbaloporetta (Millettiana) millettii 32, 33
saundersi 180, 181 CYMBALOPORIDAE 17, 32, 33
salida 178, 181*, 190, 191
'Blowiella' bentonensis 213 DISCORBIDAE 17,32,33
Bolliella 232, 233
Favusella 3, 4, 24, 25, 30, 54, 55, 57, 58, 59, 61-3,
Candeina 231 85, 108, 229
nitida 231, 242, 243 aft. washitensis 61
CANDEINIDAE 3, 4, 230, 235, 242, 243 confusa 59, 60, 68, 69
CANDEININAE 230, 231 hedbergellaeformis 61, 62, 76, 77
Cassigerinella 53, 121, 238 hiltermanni 58, 60, 76, 77
Catapsydrax 20, 30, 57 hoterivica 40, 41, 54, 58,60*,61,62,64,65,
Caucasella 85 66, 67
handousi 107 nitida 55, 61, 62, 72, 73
hoterivica 87, 88 orbiculata 59, 68, 69
Clavatorella 233 papagayosensis 62, 74, 75
Claviblowiella 56, 82, 83,177,178, 181, 182, 186, pessagnoi 61, 72, 73
237 planata 59
minai 178,182*,192,193 quadrata 62
saundersi 56, 178, 180, 181 *, 190, 191, scitula 62
192,193 sp.59
sigali 178, 182, 192, 193 stiftia 58, 61*, 63, 68, 69
Clavihedbergella 103, 163, 166 voloshinae 62
bizonae 164 washitensis 4, 8, 9, 57, 58, 59*, 60, 61, 63, 68, 69,
bollii 164 70, 71
eocretacea 55, 129, 163, 164 wenoensis 59, 74, 75
 SYSTEMATIC INDEX 261

FAVUSELLACEA 3-5,17,19,20,21,29,53,54, Globigerinelloides 56, 83,177,179,182,186,211,

58, 59, 63, 85, 229, 230, 238-40 213, 239, 245
FAVUSELLIDAE 29, 30, 54, 58, 59, 62, 236 algeriana 182
algerian us 56, 178, 182*, 183, 194, 195, 215
Globanomalina praepumilio 234 aptiense 212
Globastica 4 asper 245
Globigerina 3, 232, 236 harri 214
aequilateralis 232 bentonensis 212, 213
almadenensis 204 blowi 179, 180
angustiumbilicata 231 bollii 212
aptica 128, 129 caseyi 212, 213
bathoniana 27 cepedai 186
bulloides 14, 15, 230, 232 cushmani 212
conica 24 eaglefordensis 213
digitata 232 ferreolensis 178, 179, 182, 183*, 196, 197
dutertrei 233 gottisi 180
gaurdakensis 20, 50, 51 macrocamerata 183, 196, 197
globigerinelloides 209 maridalensis 180
helvetojurassica 27, 28, 29 manai 182
hoterivica 27, 60, 62, 85, 88 saundersi 181
infracretacea 124, 128 sigali 182
infracretacea gargasiana 107, 128 sp. (Ben Haj Ali, 1987) 212
jurassica 25 GLOBIGERINIDAE 229-31, 239
kugleri 87 GLOBIGERININA 1-5, 17,20,54,62,63,121,
oxfordiana 26 215,238
quadricamerata 60, 61, 64, 65 GLOBIGERININAE 231, 232, 236
roblesae 165 Globigerinita 231
rubra 232 iota 231
siphonifer 232 naparimaensis 231
sp. (Balakhmatova, 1953) 23 GLOBIGERINITINAE 230, 231, 235, 236, 241,
sp. D9 (Hecht, 1938) 125 242,243
sp. (Obreg6n de la Parra, 1959) 106 Globigerinoides 232
spuriensis 20, 50, 51 Globoconusa 4
tardita 60, 61, 64, 65 daubjergensis 20, 50, 51
terquemi 26 Globorotalia 234
tuschepsensis 55, 105, 109 cultrata gr. 234
washitensis 55, 59 gemma 231
'Globigerina' avarica 22 opima opima 233
balakhmatovae 23 GLOBOROTALIINAE 231, 233
bathoniana 27 Globotruncana 30, 239
Globigerina (Conoglobigerina) avarica 22 GLOBOTRUNCANIDAE 3, 204, 239
dagestanica 20 Globuligerina 3, 19, 20,21, 26, 28, 29, 53, 54, 57,
Globigerina (Eoglobigerina) balakhmatovae 20, 23, 62
38, 39 araksi 22, 36, 37
'Globigerina helvetojurassica' 28 balakhmatovae 23
Globigerina infracretacea 130 bathoniana 4, 5, 19,21,27*,46,47,229
GLOBIGERINACEA 2-5, 30, 53, 55, 83, 85, 104, calloviensis 21, 27*, 28, 40, 41
124,163,179,203,205,211,229,230,236, caucasica 24
238-42 compressa 53, 88
Globigerinatella 3 geczyi 20, 50, 51
Globigerinella 233 gulekhensis 24, 25
gottisi 180 hauterivica 60
262 SYSTEMATIC INDEX

hoterivica 60, 87 delrioensis 128

meganomica 25 excelsa 126
mejezensis 53, 54, 88, 100, 101 globigerinelloides 209
neili 53, 54, 88 gorbachikae 127
oxfordiana 4--6, 7, 8, 9, 19,21,26*,27,28,48, hiltermanni 60
49,229 hispaniae 128
parva 29 hoterivica 61
quadricamerata 60 infracretacea 124, 125, 128, 129
spiralis 53, 54, 88, 100, 101 infracretacea gargasiana 107
stellapolaris 29 intermedia 209
tardita 60 kuhryi 165
Glogigerinelloides bizonae 164 lobocaensis 165
Gorbachikella 53-5, 63, 82, 83, 85, 86, 88, 89, 103, maslakovae 163, 165, 168, 169
237 modesta 209
anteroapertura 85, 86, 87*, 94,95 occulta 130
depressa 54, 86, 87, 88*, 96, 97, 100,101 planispira 132, 204, 206, 207
grandiapertura 54, 85, 86, 87, 88*, 98, 99, 100, 101 praetrochoidea 130, 131
kugleri 55, 86,87*,88,90,91,92,93 praetrocoidea 130
neili 54, 86, 87, 88*, 100, 101 pseudotrocoidea 133
sp. 92, 93 punctata 133, 209
Gubkinella bathoniana 27 roblesae 165
Guembelitria 53, 121, 238 sigali 108
similis 163, 165, 168, 169
Haeuslerina 20, 21, 28-30, 59 trocoidea 4, 6, 7, 133, 134,203*,204-6,207
helvetojurassica 4,21,28*,29,42,43 washitensis 59
parva 21, 29*, 40, 41 HETEROHELICACEA 3, 53, 238
Hantkenina 185
multispina 179 Jurassorotalia 19
Hastigerina 232, 236
murrayi 232 Leupoldina 56, 82, 83,177,178,181,184-6,222,
pelagica 12, 13 237
Hastigerina (Bolliella) adamsi 232 cabri 184
H astigerinella gr. cabri 184, 185, 198, 199
bermudezi 233 pentagonalis 184
bizonae 164 protuberans 56, 178, 184*, 198, 199
riedeli 232 pustulans 184
Hastigerinelloides cepedai 186 reicheli 184
HASTIGERININAE 231, 235, 236 Lilliputianella 55, 82, 83, 123, 133, 161, 162, 163,
Hastigerinopsis 231, 232, 236 166,236,237
digitiformans 231 bizonae 162,164*,172,173
Haynesina 240 eocretacea 163, 164
Hedbergella (Hedbergella) sigali 108, 109 globulifera 133,162,163,164*,165,168,169
Hedbergella hoterivica 107 kuhryi162, 164, 165*, 170, 171
HEDBERGELLIDAE 30, 53, 55, 56, 83, 121, 133, labocaensis 162,163,165*,170,171
182,203,204,211,236,239-41,245,246 longorii 55, 162,163,164*, 170, 171
Herbergella 3, 19,56,83, 103, 107, 121, 125, 131, roblesae 162, 164, 165*, 172, 173
133,166,203,205,209,236,239,245,246 sp. 1 (BouDagher-Fadel, 1995) 163
aptiana 125 Lilliputianelloides 55, 83, 161,162,163,166
aptica 125, 128, 129 eocretaceus 162,163*, 168, 169
bollii 164, 172, 173 Loeblichella
cf. occulta 130 convexa 126
cf. sigali 109 moulladei 126, 142, 143
 SYSTEMATIC INDEX 263

Mariannenina 19 107, 109, 110, 121, 124, 132, 161, 163, 166,
177,237,238
Neoconorbina (Tretomphaloides) concinna grigelisae 106
32, 33 grigelisi perforare 106
Neogloboquadrina 233, 234 handousi104, 105, 107*, 114, 115
pachyderma 29 pseudosigali 104,105, 107*, 114, 115
Nonionina pelagica 232 ruka s.s. 104, 105, 107*, 110, 114, 115
ruka contritus 104, 105, 107*, 108, 110, 114, 115
Oberhauserella 19 ruka pap illata 104, 105, 108*, 110, 116, 117, 133
quadrilobata 19, 34, 35 sigali s.s. 104,105,108*, 116, 117, 118, 119
OBERHAUSERELLIDAE 19 sigali compacta 104, 105, 109*, 116, 117
Orbulina 3, 233 sigali rasilis 108
universa 12, 13, 233 tatianae 79, 104, 105, 109*, 118, 119
ORBULININAE 231, 233, 235 tuschepsensis s.s. 103,104, 105*, 106, 107, 109,
arcadia 3, 232 112, 113, 132
riedeli 14, 15,230 tuschepsensis grigelisi 104, 105, 106*, 112, 113
tuschepsensis perforare 104, 105, 106*, 112, 113,
Paragloborotalia 30, 121, 233, 234 133
incompta 234 yakovlevae 104,105,109*,118,119
Parasubbotina 3 PRAEHEDBERGELLIDAE 3, 4, 30, 53, 55, 63,
Phanerostomum asperum 245 79,81,82,83,85,89,105,133,163,166,177,
Planohedbergella 211, 214, 245, 246 185, 204, 211, 229, 235-42
ehrenbergi 245*, 248, 249 Praemurica 20, 50, 51
Planomalina 3, 56, 83, 209, 210, 211, 214, 215, Praeorbulina 3
246 Pseudo textularia 238
apsidostroba 214 Pullenia sphaeroides var. obliquiloculata 233
blowi 55, 179, 180 Pulleniatina 233
buxtorji 56, 183,210,214*,215,226,227,246 PULLENIATININAE 231, 233
caseyi 212, 220, 221, 222, 223 Pulvinulina menardii var. tumida 233
cheniourensis 183, 210, 215*, 226, 227 Pylodexia uvula 14, 15, 242, 243
ehrenbergi 245
escheri 181 Reticuloglobigerina 59, 60
maridalensis 180 washitensis 59
praebuxtorji 213 ROBERTININA 19
saundersi 181 Rosalina 240
Planomalina (Globigerinelloides) bentonensis 213 ROTALIINA 239, 240
caseyi 212 Rotalipora 30, 204, 236, 240
PLANOMALINIDAE 53, 55, 82, 83, 182, 186, Rugoglobigerina 3
204, 211, 239-41 rugosa 10, 11
Planulina
buxtorji 214 Schackoina 56, 82, 83, 177, 184-6, 213, 222, 237
cheniourensis 215 cabri 178, 184, 185*, 198, 199
Praeglobotruncana 3, 56, 121, 204, 235, 246 cenomana 178,184*,200,201
delrioensis 204, 206, 207 cenomana pentagonalis 186
hessi hessi 126 cepedai178, 186*, 200, 201
modesta 209 muitispina 179
rohri 204 pentagonalis 178, 185, 186*, 200, 201
stephani 204, 206, 207 pustulans pustulans 184
Praegubkinella 19 pustulans quinquecamerata 184
fuchsi 34, 35 reich eli 184, 186, 200, 201
ramosa 19, 34, 35 SCHACKOINIDAE 4, 53, 55, 82, 83, 110, 179,
Praehedbergella 55, 63, 82, 83, 85, 103, 104, 105, 181, 185, 186, 236, 239-41
264 SYSTEMATIC INDEX

Schmidita 19 roberti 134, 203, 205

Siderolina cenomana 56, 184 Tinophodella 231
Siderolites 185 ambitacrena 231
Sphaeroidina bulloides excavata 232 glutinata 230, 242, 243
Sphaeroidinella 232 Truncatulina humilis 234
SPHAEROIDINELLINAE 231, 232 Turborotalita 234
TURBOROTALITINAE 231, 234
Tenuitella 121, 231
anfracta 242, 243 Woletzina 20
Tenuitellinata 231 jurassica 25
bradyi 14, 15,230,242,243 Wondersella 55, 82, 123, 161, 162, 166,
Tenuitellita 231 238
Thalmanninella 204 athersuchi 55,80,162,166*,
Ticinella 203, 205, 240 174, 175
 Subject Index

Abu Dhabi (U.A.E.) 106-10, 112, 114, 116, 118, 121, 124-33, 136,
Shuaiba Formation 166, 174, 237 138, 140, 142, 144, 146, 148, 150, 152, 154,
Zakum-1 Well 174 156,158,161,164,165,168,170,172,174,
Accessory apertures 58, 59, 203, 204, 232, 234 177,179-86,188,190,192,194,196,198,200,
Albian 2, 6, 55-7, 59-62, 68, 72, 76,125,127,128, 203-6,209,211,213-15,218,220,222,226,
132, 133, 136, 148, 158, 166, 180, 181, 184-6, 229,235-7,240,241,245
192,203-5,209,211-16,222,224,226,235, Aragonite 1,4,5,8,9,17,19,27,54,62
240, 245 Atherfield Clay 108
Algeria 4, 183, 215 Azerbaijan 2,6,22,24,25,27,61,63,126-8,179
Jebel Menaouer 194 Gulekh 42
Oued Cheniour section 226 Nakhichevan 23, 44, 46
A. mayaroensis Zone 10 Nardaran 6,138,140
Ammonites 2, 86, 215 Negramskoye Gorge 36, 44, 46
Angola Basin 179 Ugakh 42
Anoxicity 205, 235, 237, 240
Aperturallip(s) 1, 22, 29, 54, 133,205,230-34 Bajocian 17-20, 22-4, 27, 36, 44, 46, 50, 240
Aperture Barremian 54-6,61,63,64,87,88,90,92,
extraumbilical 105, 235 94,96,98,103,106-10,112,114,116,118,
interiomarginal 55, 163, 209, 211, 213, 230-33, 121, 123, 125, 126, 129-31, 150, 154, 156,
245,246 161, 163, 165, 166, 168, 177, 179, 184, 186,
intra-extraumbilical 20, 26, 28-30, 40-3, 54, 55, 214,237,240,241
59,61,62,72,73,82,85,107,125,126,163, Bathonian 19-20, 22-5, 27, 28, 30, 38, 44, 46, 50
231 Bavaria 125, 127, 130
intraumbilica120, 22, 24-6, 30, 42, 43, 54, 55, 57, Bavarian Calcareous Alps 204
59,61,63,82,85,87-95,98,99,105,231-3, Bermuda 12
242,243 Berriasian 18, 24, 25, 30, 42, 57, 60, 61, 63, 66
loop-shaped 20, 26-8, 40, 41, 46-9 B. gorbachikae Zone 179, 183
umbilical-extraumbilical 55, 164, 233 Biology 1, 230-6, 238-40
Aptian 3, 6, 53, 55, 56, 60-4, 72, 85, 87-9, 103, Birmensdorfer Schichten 28, 42
266 SUBJECT INDEX

Bulgaria 24, 26 Egypt, Sinai 126, 131, 164, 165, 184, 204
Strubel 18 England
Bulla(e) 22, 23, 25-7, 30, 46, 48, 49, 54, 57, 60, 61, Arlesey brickpit 6, 213, 220, 222
63, 66-71, 231, 240 Atherfield Clay 108
Gault Clay 6, 213, 222
Calcite 1, 4, 5, 12, 13, 53, 55, 232, 234, 235, Isle of Wight 108, 248
238,245 North Sea wells 87, 103, 106-9, 112, 114, 116,
California 87, 204 118, 125-7, 129-31, 144, 148, 150, 152, 154,
Callovian 19,25,26, 28, 40, 44, 46 156, 158, 237
Campanian 211, 212 Speeton Cliff, Yorkshire 6, 103, 106, 109, 118,
Canada 124, 129, 130, 132, 136, 138, 148, 150, 156,
Grand Banks, Newfoundland 27 164, 165, 168, 170, 180, 181, 186, 188, 190,
Scotia Shelf Offshore wells 19, 27, 61, 62, 64, 66, 198, 236
70 Eocene 3, 185
Canal systems 1 Ethiopia 27
Canaliculate wall 6, 8, 9, 66, 67
Caucasus 23-5, 40, 42, 61, 62, 64, 128 Falciferum Zone (Toarcian) 34
Ilkaya 148 Favose reticulations 3, 20, 24, 25, 30, 54, 59-67, 70,
Kura River 64 71
Tuscheps River 64, 106, 112, 132, 158 Floatation structures 17, 32, 33
Cenomanian 3, 8, 55-7, 59, 63, 70, 76, 166, 185, France 4, 23, 27, 109, 127, 128, 164, 165, 180, 183,
186,200,206,209,211,213-15,236,239, 204, 212, 214, 218
240, 246 Aube 125
Coiling Basses-Alpes 108
planispiral 55, 82, 110, 132, 177, 181-6, 194, Hautes-Alpes 108, 116, 118, 165
211-15,230,231,245,246 Jura 23, 24
streptospiral82, 123, 166, 174, 175,230,231,233 La Bedoule 165, 172
trochospiral14, 54, 55, 82, 105, 121, 127, 163, La Drome 108, 126-8, 132, 142, 144, 164, 172,
166,185,230-2,238,246 179, 180, 183, 196, 236
Coniacian 185, 248 Le Havre 27, 48
Cortex 230-2, 234 Pierre Fiche (Provence) 188
Costellae 3, 10, 11 Vaucluse 109
Cretaceous 2-4, 19, 20, 53, 85, 88, 89, 103, 166,
177,184,186,204,205,211,212,214,235, Gametes 17, 18, 32, 232
237, 238, 242, 245 Gargasian (Late Aptian) 109, 165, 180, 182-4, 188,
Crimea 4, 24-9, 40, 44, 46, 64, 108, 127, 128, 131, 196, 200, 204, 212, 218
164-6,168,181,183,184,204 Gault Clay 6, 213, 222
Kacha River 2,118, 126-8, 131, 144, 146, 152, Germany 27,130, 185,200
154, 156, 176, 179-81, 183, 184, 186, 188 Bavaria 125, 127, 130
Krasnaya 6,126,140,168,172,181,190,194 Island of Rugen 245
Predoroshnaya 125, 136 Luneburg 185
Simferopol 6, 206 Mittelland-Kanal, Braunschweig 138
Cytoplasm 1,3,205,232,235,240 Moltzow, Mecklenburg 185
extrathalamous 1, 3, 17, 30, 205, 211, 234-6, Otto Gott pit, Sarstedt 148
238-40 Wiechendorf 1/86 Borehole 128, 164, 179-81,
intrathalamous 205, 235, 236 184
Zeltberg brickworks, near Hamburg 60, 76
Dagestan 4, 22-4, 36, 38 G. ferreolensis Zone 109, 179, 180, 212, 218
Danian 4, 20, 55, 103, 121
Diploid 18 Haploid 18
DSDP Site 397 (N.E. Atlantic) 61, 68, 179 Hauterivian 25, 40, 55, 61-4, 68, 87, 88, 90, 92, 103,
Dysoxicity 80, 166, 205, 236, 237 107,108,110,114,179,229,235,237,240,241
 SUBJECT INDEX 267

Hettangian 50 North Sea wells 87, 103, 106-9, 112, 114, 116, 118,
Holocene 238 125-7, 129-31, 144, 148, 150, 152, 154, 156,
Holoplanktonic 17-19, 53, 57, 229, 240 158,237
Hungary, Gerecse Mountains 50
Hyaline 4 Oligocene 121
Oxfordian 4, 6, 8, 25, 27-9, 42, 48
Indian Ocean 12, 214, 224, 226
Israel 166 Palaeobathymetry 237, 240
Italy, Rocchetta section 50 Palaeocene 3, 50
Palaeoecology 2,17-19,59,80,84,131,161,186,
Jurassic 2, 3, 5, 19, 20, 23, 27, 30, 50, 53, 203-5,229,234-41
57, 235 Perforation cones 3, 6, 7, 55, 79, 80, 105-8, 112,
113, 121, 124, 126, 128, 131, 133, 138-41, 148,
Kimmeridgian 27, 29, 40
149, 182, 194-7, 211, 230, 235, 238, 239, 241
Lithuania 27 Perforations/perforate 1-5, 17, 59, 148,205, 230-3,
L. protuberans Zone 179 235,239
Peripheral keel 56, 183, 233, 239
Maastrichtian 10, 56, 121, 177, 186,211 Pillars 1
Macroperforations/macroperforate 6, 7, 10, 11, 14, Planispira155, 82, 110, 132, 177, 181-6, 194,
15, 17, 30, 50, 51, 55, 56, 82, 103, 121, 133, 211-15, 230, 231, 245, 246
183,184,186,203-7,209,211-23,226,230, Plugs 1
231, 235, 236, 245, 246, 248 249 Poland 4, 27, 46
M. coranguinum Zone 248 Ogrodzieniec 46
Megalospheric (generation) 18, 19, 22, 23, 25, 26 Pores 2, 3, 12, 13, 216, 217, 241
Meiosis 18 Porticus(i) 1, 55, 56, 106, 108, 109, 112, 124-6, 129,
Meroplanktonic 18, 22, 229 132,133,144,148,164, 179, 181, 182, 194,
Mexico 63, 68, 87, 103, 127, 130, 132, 166, 180, 181, 195,203-5,211-13,240,245
183, 185, 186, 204, 214 Portugal 27
La Perra Formation (La Boca Canyon) 128, 146, Pseudomuricae 3, 6, 7-9, 20, 23-9, 36-9, 42, 43, 46,
150,164-6,168, 170, 172, 179, 182, 192,200, 47,54,57,59,62
218 Pseudopodia 3, 17, 18, 205, 235, 239
Sabinas Basin 59, 62 Pustules 3, 4, 14, 15,230, 235, 242, 243
Sierra de Papagayos 60, 62, 74, 76
Microperforations/microperforate 4, 6-9, 14, 15, Recent 12, 14, 242
36-9, 46, 47, 53-6, 59, 63, 66, 67, 70, 71, 79, Romania 204
82, 85, 89, 94-7, 103, 105, 112, 118, 119, 121, Izvorul Valley 163, 168
131, 136-9, 163, 181, 182, 184-6, 190, 191, Russia 4
194-7, 203, 204, 209, 211, 213, 230, 235, 236, Moscow Region 4, 6, 27
242,243 Pechora River Basin 29, 44, 46
Microspheric (generation) 18, 19,22-5 Upper Volga Basin 27,48
Miocene 232 Russian Plate 4
Mitosis 18
Multiple apertures 31, 233 Santonian 248
Muricae/muricate 3, 6, 7, 10,11,28, 50, 51, 55, 56, S. cabri Zone 106, 109, 112, 118, 126, 130, 136,
103, 121, 133, 186, 203-7, 209, 213, 214, 216, 138, 142, 146, 148, 150, 156, 164, 165, 168,
217,224-7,239-41,245,248,249 170,172,179-81,183,188,190,192,198,205
Muricocarina(e) 3, 56, 121, 204, 209, 211, 213-15, Sicily 185
224, 225, 239, 246 Sinemurian 50
Spain 127, 166, 186, 214, 215
Nannofossils 2, 61, 86, 179, 205 Argos Formation 128, 170, 200
Newfoundland (Grand Banks) 27 EI Burrueco section 224
North Africa 85, 88, 161 Spines/spinose 3, 4, 12-15, 230-33, 235, 236
268 SUBJECT INDEX

Streptospiral82, 123, 166, 174, 175, 230, 231, 233 Ukraine 4, 6, 25, 28 see also Crimea
Swiss Alps (Teysachaux) 34 U.S.A. see California, Texas, Wyoming
Switzerland 27
Birmensdorfer Schichten 28, 42 Valanginian 18, 19,24, 25, 27, 30, 53-5, 57, 61, 63,
Breggia River 204, 206, 215 64, 66, 82, 85, 88, 92, 100, 240
Eisengraben section 28, 42 Volgian 29, 44, 46
Renggeri Tonen 28
Symbionts 231-4, 239 Walvis Ridge 103
West Carpathians (Magura Flysch) 87, 88, 125-9,
Tanzania, Lindi River 215, 226 142
Tethys 17, 59, 131, 166, 179,203,211,237,240 Wyoming 236
Texas 4, 8, 62, 70, 236
Del Rio Formation 60 X-ray analysis 4, 27
Denton Formation 62
Duck Creek Formation 62, 72, 74
Index to depositories: The specimens illustrated in
Forth Worth Formation 62, 72
the plates of this book are deposited in the
Grayson Formation 60, 206
following museums:
Main Street Limestone 214
Shoal Creek 8, 60 Baku, Azerbaijan (Institute of Geology,
Washita Group 60, 62, 68, 70, 72 Azerbaijan Academy of Sciences) 36
Tithonian 18, 24, 26 Basle, Switzerland (Natural History Museum) 198
Toarcian 19, 34 Bedford, Canada (Atlantic Geoscience Centre,
Toothplates 1 Geological Survey of Canada) 42, 72
Transversarium Zone 28 Bratislava, Slovakia (Dionys Stur Institute of
Triassic 19 Geology) 114
Trinidad 87, 181, 185 Brno, Czech Republic (Czech Geological Survey)
Cuche Formation 179, 188 142
Guayaguayare 10 Bucharest, Romania (Laboratory of
Maridale Formation 180, 181, 188, 192, 204, 209, Palaeontology, University of Bucharest) 168
214,218 Budapest, Hungary (Hungarian Natural History
Montesserat-1 Well 92 Museum) 50
Piparo River 198
Trochospiral 14, 54, 55, 82, 105, 121, 127, 163, 166, Frankfurt-am-Main, Germany (Senckenberg
185, 230-2, 238, 246 Museum) 50, 138
Tubulospines 56, 177, 184-6, 198-201, 213, 222,
223, 236 Geneva, Switzerland (Museum d'Histoire
Tunisia 2, 54, 87-9, 107, 108, 126, 127, 132, 163, Naturelle de Geneve) 34
165,179,180,183-5,186,204,212,213,237
Beauvoir-I Well 90, 92, 94, 96, 98, 100, 168 Kiel, Germany (Geological-Palaeontological
Beauvoir-III Well 6, 216 Institute and Museum, University of Kiel) 60
Beauvoir-IV Well 196, 200, 222 Krasnodar, Russia (All-Union Oil and Gas
Beauvoir-VI Well 158 Geological-Prospecting Institute) 64
Jebel Chenanrafa 146, 181, 190, 198,222
Jebel Oust wells 53, 54, 92, 96, 100, 114, 200, London, England (The Natural History Museum,
226 formerly British Museum (Natural History
Jebel Rhazouane 185, 198 4,6,8,10,14,20,32,44,46,48,70,90,92,94,
Oued Guetta 126 96, 98, 109, 112, 114, 116, 118, 127, 131, 136,
Turkey 183, 184, 204 138, 140, 144, 146, 150, 152, 154, 156, 158,
Turkmenia 22-24, 50 168,170, 172, 174, 186, 188, 190, 194, 198,
Toronian 185 200,216,220,222,224,226,242,248
Lyon, France (Department of Earth Sciences,
U.A.E. see Abu Dhabi University of Lyon) 116, 182, 196
 SUBJECT INDEX 269

Moscow, Russia (Geological Faculty, Moscow St Petersburg, Russia (All-Union Petroleum

State University) 42, 44, 46, 48, 105, Scientific Research-Prospecting Institute
106, 118, 132, 152, 158, 168, (VNIGRI 40, 64,103,112
190,206
Moscow, Russia (Geological Institute, Tunis, Tunisia (National Geological Survey of
Academy of Sciences of Russia) Tunisia) 92, 100
36,38,40
Utrecht, The Netherlands (Utrecht
Paris, France (Laboratoire de Micropalt!ontologie, Micropalaeontological Collections, State
Universite de Paris) 172, 188 University of Utrecht) 224
Paris, France (Museum National d'Histoire
Naturelle) 206 Washington, D.C., U.S.A. (United States
National Museum of Natural History)
St Petersburg, Russia (F.N. Chemyshev Museum, 68,70,72,74,76,92,142,144,146,
All-Union Scientific Research Geological 148, 150, 168, 170, 172, 186, 188,
Institute) 50 192,194,196,200,218